Theropoda

Theropoda (/θɪəˈrɒpədə/ or /ˌθɪərəˈpoʊdə/,[2] from Greek θηρίον "wild beast" and πούς, ποδός "foot") or theropods (/ˈθɪərəˌpɒdz/[3][4]) are a dinosaur suborder that is characterized by hollow bones and three-toed limbs. They are generally classed as a group of saurischian dinosaurs, although a 2017 paper has instead placed them in the proposed clade Ornithoscelida as the closest relatives of the Ornithischia.[5] Theropods were ancestrally carnivorous, although a number of theropod groups evolved to become herbivores, omnivores, piscivores, and insectivores. Theropods first appeared during the Carnian age of the late Triassic period 231.4 million years ago (Ma)[6] and included the sole large terrestrial carnivores from the Early Jurassic until at least the close of the Cretaceous, about 66 Ma. In the Jurassic, birds evolved from small specialized coelurosaurian theropods, and are today represented by about 10,500 living species.

Theropods
Temporal range:
Late TriassicPresent, 231.4–0 Ma
Coelophysis bauri mount
Mounted skeleton of Coelophysis bauri, Cleveland Museum of Natural History
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Marsh, 1881
Subgroups[1]

Biology

Diet and teeth

Jinfengopteryx elegans 2
Specimen of the troodontid Jinfengopteryx elegans, with seeds preserved in the stomach region

Theropods exhibit a wide range of diets, from insectivores to herbivores and carnivores. Strict carnivory has always been considered the ancestral diet for theropods as a group, and a wider variety of diets was historically considered a characteristic exclusive to the avian theropods (birds). However, discoveries in the late 20th and early 21st centuries showed that a variety of diets existed even in more basal lineages.[7] All early finds of theropod fossils showed them to be primarily carnivorous. Fossilized specimens of early theropods known to scientists in the 19th and early 20th centuries all possessed sharp teeth with serrated edges for cutting flesh, and some specimens even showed direct evidence of predatory behavior. For example, a Compsognathus longipes fossil was found with a lizard in its stomach, and a Velociraptor mongoliensis specimen was found locked in combat with a Protoceratops andrewsi (a type of ornithischian dinosaur).

The first confirmed non-carnivorous fossil theropods found were the therizinosaurs, originally known as segnosaurs. First thought to be prosauropods, these enigmatic dinosaurs were later proven to be highly specialized, herbivorous theropods. Therizinosaurs possessed large abdomens for processing plant food, and small heads with beaks and leaf-shaped teeth. Further study of maniraptoran theropods and their relationships showed that therizinosaurs were not the only early members of this group to abandon carnivory. Several other lineages of early maniraptors show adaptations for an omnivorous diet, including seed-eating (some troodontids) and insect-eating (many avialans and alvarezsaurs). Oviraptorosaurs, ornithomimosaurs and advanced troodontids were likely omnivorous as well, and some early theropods (such as Masiakasaurus knopfleri and the spinosaurids) appear to have specialized in catching fish.[8][9]

Diet is largely deduced by the tooth morphology,[10] tooth marks on bones of the prey, and gut contents. Some theropods, such as Baryonyx, Lourinhanosaurus, ornithomimosaurs, and birds, are known to use gastroliths, or gizzard-stones.

The majority of theropod teeth are blade-like, with serration on the edges,[11] called ziphodont. Others are pachydont or phyllodont depending on the shape of the tooth or denticles. The morphology of the teeth is distinct enough to tell the major families apart,[12] which indicate different diet strategies. An investigation in July 2015 discovered that what appeared to be "cracks" in their teeth were actually folds that helped to prevent tooth breakage by strengthening individual serrations as they attacked their prey.[13] The folds helped the teeth stay in place longer, especially as theropods evolved into larger sizes and had more force in their bite.[14][15]

Skin, scales and feathers

Anchiornis feathers
Fossil of a Anchiornis, showing large preserved feather imprints

Mesozoic theropods were also very diverse in terms of skin texture and covering. Feathers or feather-like structures are attested in most lineages of theropods. (See feathered dinosaur). However, outside the coelurosaurs, feathers may have been confined to the young, smaller species, or limited parts of the animal. Many larger theropods had skin covered in small, bumpy scales. In some species, these were interspersed with larger scales with bony cores, or osteoderms. This type of skin is best known in the ceratosaur Carnotaurus, which has been preserved with extensive skin impressions.[16]

The coelurosaur lineages most distant from birds had feathers that were relatively short and composed of simple, possibly branching filaments.[17] Simple filaments are also seen in therizinosaurs, which also possessed large, stiffened "quill"-like feathers. More fully feathered theropods, such as dromaeosaurs, usually retain scales only on the feet. Some species may have mixed feathers elsewhere on the body as well. Scansoriopteryx preserved scales near the underside of the tail,[18] and Juravenator may have been predominantly scaly with some simple filaments interspersed.[19] On the other hand, some theropods were completely covered with feathers, such as the troodontid Anchiornis, which even had feathers on the feet and toes.[20]

Size

Longest theropods
Size comparison of selected giant theropod dinosaurs

Tyrannosaurus was for many decades the largest known theropod and best-known to the general public. Since its discovery, however, a number of other giant carnivorous dinosaurs have been described, including Spinosaurus, Carcharodontosaurus, and Giganotosaurus.[21] The original Spinosaurus specimens (as well as newer fossils described in 2006) support the idea that Spinosaurus is longer than Tyrannosaurus, showing that Spinosaurus was possibly 3 meters longer than Tyrannosaurus though Tyrannosaurus could still be taller and more massive than Spinosaurus.[22] Specimens of Tyrannosaurus such as Sue and Scotty are estimated to be the most massive theropods known to science. There is still no clear explanation for exactly why these animals grew so much larger than the land predators that came before and after them.

The largest extant theropod is the common ostrich, up to 2.74 m (9 ft) tall and weighing between 63.5 and 145.15 kg (140 - 320 lb).[23]

The smallest non-avialan theropod known from adult specimens is the troodontid Anchiornis huxleyi, at 110 grams in weight and 34 centimeters (1 ft) in length.[20] When modern birds are included, the bee hummingbird Mellisuga helenae is smallest at 1.9 g and 5.5 cm (2.2 in) long.[24]

Recent theories propose that theropod body size shrank continuously over a period of 50 million years, from an average of 163 kilograms (359 lb) down to 0.8 kilograms (1.8 lb), eventually evolving into modern birds. This was based on evidence that theropods were the only dinosaurs to get continuously smaller, and that their skeletons changed four times as fast as those of other dinosaur species.[25][26]

Stance and gait

Struthio camelus - Etosha 2014 (1)
An ostrich walking on a road in Etosha National Park, Namibia

As a hugely diverse group of animals, the posture adopted by theropods likely varied considerably between various lineages through time.[27] All known theropods are known to be bipedal, with the forelimbs reduced in length and specialized for a wide variety of tasks (see below). In modern birds, the body is typically held in a somewhat upright position, with the upper leg (femur) held parallel to the spine and with the forward force of locomotion generated at the knee. Scientists are not certain how far back in the theropod family tree this type of posture and locomotion extends.[27]

Non-avian theropods were first recognized as bipedal during the 19th century, before their relationship to birds was widely accepted. During this period, theropods such as carnosaurs and tyrannosaurids were thought to have walked with vertical femurs and spines in an upright, nearly erect posture, using their long, muscular tails as additional support in a kangaroo-like tripodal stance.[27] Beginning in the 1970s, biomechanical studies of extinct giant theropods cast doubt on this interpretation. Studies of limb bone articulation and the relative absence of trackway evidence for tail dragging suggested that, when walking, the giant, long-tailed theropods would have adopted a more horizontal posture with the tail held parallel to the ground.[27][28] However, the orientation of the legs in these species while walking remains controversial. Some studies support a traditional vertically oriented femur, at least in the largest long-tailed theropods,[28] while others suggest that the knee was normally strongly flexed in all theropods while walking, even giants like the tyrannosaurids.[29][30] It is likely that a wide range of body postures, stances, and gaits existed in the many extinct theropod groups.[27][31]

Nervous system and senses

Although rare, complete casts of theropod endocrania are known from fossils. Theropod endocrania can also be reconstructed from preserved brain cases without damaging valuable specimens by using a computed tomography scan and 3D reconstruction software. These finds are of evolutionary significance because they help document the emergence of the neurology of modern birds from that of earlier reptiles. An increase in the proportion of the brain occupied by the cerebrum seems to have occurred with the advent of the Coelurosauria and "continued throughout the evolution of maniraptorans and early birds."[32]

Forelimb morphology

Mummified precocial bird wings in mid-Cretaceous Burmese amber (2016) fig. 1
Mummified enantiornithean wing from Cretaceous amber

Shortened forelimbs in relation to hind legs was a common trait among theropods, most notably in the abelisaurids (such as Carnotaurus) and the tyrannosaurids (such as Tyrannosaurus). This trait was, however, not universal: spinosaurids had well developed forelimbs, as did many coelurosaurs. The relatively robust forelimbs of one genus, Xuanhanosaurus, led Dong Zhiming to suggest that the animal might have been quadrupedal.[33] However, this is no longer thought to be likely.[34]

The hands are also very different among the different groups. The most common form among non-avian theropods is an appendage consisting of three fingers; the digits I, II and III (or possibly II, III and IV),[35] with sharp claws. Some basal theropods (e.g. Herrerasaurus, Eoraptor) had four digits, and also a reduced metacarpal V. Ceratosaurians usually had four digits, while most tetanurans had three.[36]

The forelimbs' scope of use is also believed to have also been different among different families. The spinosaurids could have used their powerful forelimbs to hold fish. Some small maniraptorans such as scansoriopterygids are believed to have used their forelimbs to climb in trees.[18] The wings of modern birds are used primarily for flight, though they are adapted for other purposes in certain groups. For example, aquatic birds such as penguins use their wings as flippers.

Forelimb movement

Archaeo-deinony hands
Diagram of Deinonychus (left) and Archaeopteryx (right) forelimbs illustrating wing-like posture

Contrary to the way theropods have often been reconstructed in art and the popular media, the range of motion of theropod forelimbs was severely limited, especially compared with the forelimb dexterity of humans and other primates.[37] Most notably, theropods and other bipedal saurischian dinosaurs (including the bipedal prosauropods) could not pronate their hands—that is, they could not rotate the forearm so that the palms faced the ground or backwards towards the legs. In humans, pronation is achieved by motion of the radius relative to the ulna (the two bones of the forearm). In saurischian dinosaurs, however, the end of the radius near the elbow was actually locked into a groove of the ulna, preventing any movement. Movement at the wrist was also limited in many species, forcing the entire forearm and hand to move as a single unit with little flexibility.[38] In theropods and prosauropods, the only way for the palm to face the ground would have been by lateral splaying of the entire forelimb, as in a bird raising its wing.[37]

In carnosaurs like Acrocanthosaurus, the hand itself retained a relatively high degree of flexibility, with mobile fingers. This was also true of more basal theropods, such as herrerasaurs and dilophosaurs. Coelurosaurs showed a shift in the use of the forearm, with greater flexibility at the shoulder allowing the arm to be raised towards the horizontal plane, and to even greater degrees in flying birds. However, in coelurosaurs, such as ornithomimosaurs and especially dromaeosaurs, the hand itself had lost most flexibility, with highly inflexible fingers. Dromaeosaurs and other maniraptorans also showed increased mobility at the wrist not seen in other theropods, thanks to the presence of a specialized half-moon shaped wrist bone (the semi-lunate carpal) that allowed the whole hand to fold backward towards the forearm in the manner of modern birds.[38]

Paleopathology

In 2001, Ralph E. Molnar published a survey of pathologies in theropod dinosaur bone. He found pathological features in 21 genera from 10 families. Pathologies were found in theropods of all body size although they were less common in fossils of small theropods, although this may be an artifact of preservation. They are very widely represented throughout the different parts of theropod anatomy. The most common sites of preserved injury and disease in theropod dinosaurs are the ribs and tail vertebrae. Despite being abundant in ribs and vertebrae, injuries seem to be "absent... or very rare" on the bodies' primary weight supporting bones like the sacrum, femur, and tibia. The lack of preserved injuries in these bones suggests that they were selected by evolution for resistance to breakage. The least common sites of preserved injury are the cranium and forelimb, with injuries occurring in about equal frequency at each site. Most pathologies preserved in theropod fossils are the remains of injuries like fractures, pits, and punctures, often likely originating with bites. Some theropod paleopathologies seem to be evidence of infections, which tended to be confined only to small regions of the animal's body. Evidence for congenital malformities have also been found in theropod remains. Such discoveries can provide information useful for understanding the evolutionary history of the processes of biological development. Unusual fusions in cranial elements or asymmetries in the same are probably evidence that one is examining the fossils of an extremely old individual rather than a diseased one.[39]

Swimming

The trackway of a swimming theropod, the first in China of the ichnogenus named Characichnos, was discovered at the Feitianshan Formation in Sichuan.[40] These new swim tracks support the hypothesis that theropods were adapted to swimming and capable of traversing moderately deep water. Dinosaur swim tracks are considered to be rare trace fossils, and are among a class of vertebrate swim tracks that also include those of pterosaurs and crocodylomorphs. The study described and analyzed four complete natural molds of theropod foot prints that are now stored at the Huaxia Dinosaur Tracks Research and Development Center (HDT). These dinosaur footprints were in fact claw marks, which suggest that this theropod was swimming near the surface of a river and just the tips of its toes and claws could touch the bottom. The tracks indicate a coordinated, left-right, left-right progression, which supports the proposition that theropods were well-coordinated swimmers.[40]

Evolutionary history

Herrerasaurusskeleton
Possible early forms Herrerasaurus (large) and Eoraptor (small)

During the late Triassic, a number of primitive proto-theropod and theropod dinosaurs existed and evolved alongside each other.

The earliest and most primitive of the theropod dinosaurs were the carnivorous Eodromaeus and the herrerasaurids of Argentina (as well as, possibly, the omnivorous Eoraptor). The herrerasaurs existed during the early late Triassic (Late Carnian to Early Norian). They were found in North America and South America and possibly also India and Southern Africa. The herrerasaurs were characterised by a mosaic of primitive and advanced features. Some paleontologists have in the past considered the herrerasaurians to be members of Theropoda, while other theorized the group to be basal saurischians, and may even have evolved prior to the saurischian-ornithischian split. Cladistic analysis following the discovery of Tawa, another Triassic dinosaur, suggests the herrerasaurs likely were early theropods.[41]

The earliest and most primitive unambiguous theropods (or alternatively, "Eutheropoda"—'True Theropods') are the Coelophysoidea. The Coelophysoidea were a group of widely distributed, lightly built and potentially gregarious animals. They included small hunters like Coelophysis and (possibly) larger predators like Dilophosaurus. These successful animals continued from the Late Carnian (early Late Triassic) through to the Toarcian (late Early Jurassic). Although in the early cladistic classifications they were included under the Ceratosauria and considered a side-branch of more advanced theropods,[42] they may have been ancestral to all other theropods (which would make them a paraphyletic group).[43][44]

The somewhat more advanced ceratosaurs (including Ceratosaurus and Carnotaurus) appeared during the Early Jurassic and continued through to the Late Jurassic in Laurasia. They competed alongside their more anatomically advanced tetanuran relatives and—in the form of the abelisaur lineage—lasted to the end of the Cretaceous in Gondwana.

The Tetanurae are more specialised again than the ceratosaurs. They are subdivided into the basal Megalosauroidea (alternately Spinosauroidea) and the more derived Avetheropoda. Megalosauridae were primarily Middle Jurassic to Early Cretaceous predators, and their spinosaurid relatives' remains are mostly from Early and Middle Cretaceous rocks. Avetheropoda, as their name indicates, were more closely related to birds and are again divided into the Allosauroidea (the diverse carcharodontosaurs) and the Coelurosauria (a very large and diverse dinosaur group including the birds).

Thus, during the late Jurassic, there were no fewer than four distinct lineages of theropods—ceratosaurs, megalosaurs, allosaurs, and coelurosaurs—preying on the abundance of small and large herbivorous dinosaurs. All four groups survived into the Cretaceous, and three of those—the ceratosaurs, coelurosaurs, and allosaurs—survived to end of the period, where they were geographically separate, the ceratosaurs and allosaurs in Gondwana, and the coelurosaurs in Laurasia.

Of all the theropod groups, the coelurosaurs were by far the most diverse. Some coelurosaur groups that flourished during the Cretaceous were the tyrannosaurids (including Tyrannosaurus), the dromaeosaurids (including Velociraptor and Deinonychus, which are remarkably similar in form to the oldest known bird, Archaeopteryx[45][46]), the bird-like troodontids and oviraptorosaurs, the ornithomimosaurs (or "ostrich dinosaurs"), the strange giant-clawed herbivorous therizinosaurs, and the avialans, which include modern birds and is the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event.[47] While the roots of these various groups are found in the Middle Jurassic, they only became abundant during the Early Cretaceous. A few paleontologists, such as Gregory S. Paul, have suggested that some or all of these advanced theropods were actually descended from flying dinosaurs or proto-birds like Archaeopteryx that lost the ability to fly and returned to a terrestrial habitat.[48]

On July 31, 2014, scientists reported details of the evolution of birds from other theropod dinosaurs.[25][49][26] Among the features linking theropod dinosaurs to birds are a furcula (wishbone), air-filled bones, brooding of the eggs, and (in coelurosaurs, at least) feathers.

Classification

History of classification

Othniel Charles Marsh - Brady-Handy
Othniel Charles Marsh, who coined the name Theropoda. Photo c. 1870

O. C. Marsh coined the name theropoda (meaning "beast feet") in 1881.[50] Marsh initially named Theropoda as a suborder to include the family Allosauridae, but later expanded its scope, re-ranking it as an order to include a wide array of "carnivorous" dinosaur families, including Megalosauridae, Compsognathidae, Ornithomimidae, Plateosauridae and Anchisauridae (now known to be herbivorous sauropodomorphs) and Hallopodidae (subsequently revealed as relatives of crocodilians). Due to the scope of Marsh's Order Theropoda, it came to replace a previous taxonomic group that Marsh's rival E. D. Cope had created in 1866 for the carnivorous dinosaurs: Goniopoda ("angled feet").[34]

By the early 20th century, some paleontologists, such as Friedrich von Huene, no longer considered carnivorous dinosaurs to have formed a natural group. Huene abandoned the name "Theropoda", instead using Harry Seeley's Order Saurischia, which Huene divided into the suborders Coelurosauria and Pachypodosauria. Huene placed most of the small theropod groups into Coelurosauria, and the large theropods and prosauropods into Pachypodosauria, which he considered ancestral to the Sauropoda (prosauropods were still thought of as carnivorous at that time, owing to the incorrect association of rauisuchian skulls and teeth with prosauropod bodies, in animals such as Teratosaurus).[34] In W. D. Matthew and Barnum Brown's 1922 description of the first known dromaeosaurid (Dromaeosaurus albertensis[51]), they became the first paleontologists to exclude prosauropods from the carnivorous dinosaurs, and attempted to revive the name "Goniopoda" for that group, but other scientists did not accept either of these suggestions.[34]

AllosaurusAMNH5753
Allosaurus was one of the first dinosaurs classified as a theropod.

In 1956, "Theropoda" came back into use—as a taxon containing the carnivorous dinosaurs and their descendants—when Alfred Romer re-classified the Order Saurischia into two suborders, Theropoda and Sauropoda. This basic division has survived into modern paleontology, with the exception of, again, the Prosauropoda, which Romer included as an infraorder of theropods. Romer also maintained a division between Coelurosauria and Carnosauria (which he also ranked as infraorders). This dichotomy was upset by the discovery of Deinonychus and Deinocheirus in 1969, neither of which could be classified easily as "carnosaurs" or "coelurosaurs". In light of these and other discoveries, by the late 1970s Rinchen Barsbold had created a new series of theropod infraorders: Coelurosauria, Deinonychosauria, Oviraptorosauria, Carnosauria, Ornithomimosauria, and Deinocheirosauria.[34]

With the advent of cladistics and phylogenetic nomenclature in the 1980s, and their development in the 1990s and 2000s, a clearer picture of theropod relationships began to emerge. Jacques Gauthier named several major theropod groups in 1986, including the clade Tetanurae for one branch of a basic theropod split with another group, the Ceratosauria. As more information about the link between dinosaurs and birds came to light, the more bird-like theropods were grouped in the clade Maniraptora (also named by Gauthier in 1986). These new developments also came with a recognition among most scientists that birds arose directly from maniraptoran theropods and, on the abandonment of ranks in cladistic classification, with the re-evaluation of birds as a subset of theropod dinosaurs that survived the Mesozoic extinctions and lived into the present.[34]

Major groups

Ceratosaurus mounted
Ceratosaurus, a ceratosaurid
Irritator challengeri mount 01
Irritator, a spinosaurid
MicroraptorGui-PaleozoologicalMuseumOfChina-May23-08
Microraptor, a dromaeosaurid
Passer domesticus male (15)
Passer domesticus, an avian, and the world's most widespread extant wild theropod.[52]

The following is a simplified classification of theropod groups based on their evolutionary relationships, and organized based on the list of Mesozoic dinosaur species provided by Holtz (2008).[53] A more detailed version can be found at Dinosaur classification. The cross (†) is used to signify groups with no living members.

Relationships

The following family tree illustrates a synthesis of the relationships of the major theropod groups based on various studies conducted in the 2010s.[54]

Theropoda

Herrerasauridae Herrerasaurus ischigualastensis Illustration

Eoraptor

Eodromaeus

Daemonosaurus

Tawa

 Neotheropoda 

 †Coelophysoidea  Coelophysis size flipped

Dilophosauridae Dilophosaurus wetherilli (flipped)

Averostra

Ceratosauria Ceratosaurus nasicornis DBCarnotaurus DB 2 white background

Tetanurae

Megalosauroidea Torvosaurus tanneri Reconstruction (Flipped)Спинозавр - новая реконструкция (flipped)

Avetheropoda

Allosauroidea Allosaurus RevisedMapusaurus BW

Coelurosauria Sinosauropteryx color Hypothetical Deinocheirus (flipped) Tyrannosaurus-rex-Profile-steveoc86 Utahraptor Restoration (flipped) Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg

A large study of early dinosaurs by Dr Matthew G. Baron, David Norman and Paul M. Barrett (2017) published in the journal Nature suggested that Theropoda is actually more closely related to Ornithischia, to which it formed the sister group within the clade Ornithoscelida. This new hypothesis also recovered Herrerasauridae as the sister group to Sauropodomorpha in the redefined Saurischia and suggested that the hypercarnivore morphologies that are observed in specimens of theropods and herrerasaurids were acquired convergently.[55][56] However, this phylogeny remains controversial and additional work is being done to clarify these relationships.

See also

References

  1. ^ Holtz, Thomas R., Jr. (2012). Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
  2. ^ See analogous pronunciation of Gastropoda - definition of Gastropoda in English from the Oxford dictionary.
  3. ^ "Theropod". Merriam-Webster Dictionary.
  4. ^ "Theropod". Dictionary.com Unabridged. Random House.
  5. ^ Baron, M.G., Norman, D.B., and Barrett, P.M. (2017). A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature, 543: 501–506. doi:10.1038/nature21700
  6. ^ Alcober, Oscar A.; Martinez, Ricardo N. (19 October 2010). "A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina". ZooKeys (63): 55–81. doi:10.3897/zookeys.63.550. PMC 3088398. PMID 21594020. [1]
  7. ^ Zanno, Lindsay E.; Gillette, David D.; Albright, L. Barry; Titus, Alan L. (25 August 2010). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution". Proceedings of the Royal Society B. 276 (1672): 3505–3511. doi:10.1098/rspb.2009.1029. PMC 2817200. PMID 19605396.
  8. ^ Longrich, Nicholas R.; Currie, Philip J. (February 2009). "Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae". Cretaceous Research. 30 (1): 239–252. doi:10.1016/j.cretres.2008.07.005.
  9. ^ Holtz, T.R.; Jr; Brinkman, D.L.; Chandler, C.L. (1998). "Dental morphometrics and a possibly omnivorous feeding habit for the theropod dinosaur Troodon". GAIA. 15: 159–166.
  10. ^ Hendrickx, C.; Mateus, O. (2014). "Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth". Zootaxa. 3759: 1–74. doi:10.11646/zootaxa.3759.1.1. PMID 24869965.
  11. ^ Hendrickx, Christophe; Mateus, Octávio; Araújo, Ricardo (2015). "A proposed terminology of theropod teeth (Dinosauria, Saurischia)". Journal of Vertebrate Paleontology (Submitted manuscript). 35 (e982797): e982797. doi:10.1080/02724634.2015.982797.
  12. ^ Hendrickx, C; Mateus, O (2014). "Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth". Zootaxa. 3759: 1–74. doi:10.11646/zootaxa.3759.1.1. PMID 24869965.
  13. ^ Geggel, Laura (28 July 2015). "One tough bite: T. rex's teeth had secret weapon". Fox News. Retrieved 1 August 2015.
  14. ^ "Special Serrations Gave Carnivorous Dinosaurs an Evolutionary Edge".
  15. ^ "Developmental and evolutionary novelty in the serrated teeth of theropod dinosaurs".
  16. ^ Bonaparte, Novas, and Coria (1990). "Carnotaurus sastrei Bonaparte, the horned, lightly built carnosaur from the Middle Cretaceous of Patagonia." Contributions in Science (Natural History Museum of Los Angeles County), 416: 41 pp.
  17. ^ Göhlich, U.B.; Chiappe, L.M. (16 March 2006). "A new carnivorous dinosaur from the Late Jurassic Solnhofen archipelago" (PDF). Nature. 440 (7082): 329–332. doi:10.1038/nature04579. PMID 16541071.
  18. ^ a b Czerkas, S.A., and Yuan, C. (2002). "An arboreal maniraptoran from northeast China." Pp. 63–95 in Czerkas, S.J. (Ed.), Feathered Dinosaurs and the Origin of Flight. The Dinosaur Museum Journal 1. The Dinosaur Museum, Blanding, U.S.A. PDF abridged version
  19. ^ Goehlich, U.B.; Tischlinger, H.; Chiappe, L.M. (2006). "Juraventaor starki (Reptilia, Theropoda) ein nuer Raubdinosaurier aus dem Oberjura der Suedlichen Frankenalb (Sueddeutschland): Skelettanatomie und Wiechteilbefunde". Archaeopteryx. 24: 1–26.
  20. ^ a b Xu, X.; Zhao, Q.; Norell, M.; Sullivan, C.; Hone, D.; Erickson, G.; Wang, X.; Han, F. & Guo, Y. (February 2009). "A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin". Chinese Science Bulletin. 54 (3): 430–435. doi:10.1007/s11434-009-0009-6. Abstract
  21. ^ Therrien, F.; Henderson, D. M. (2007). "My theropod is bigger than yours...or not: estimating body size from skull length in theropods". Journal of Vertebrate Paleontology. 27 (1): 108–115. doi:10.1671/0272-4634(2007)27[108:MTIBTY]2.0.CO;2.
  22. ^ dal Sasso, C.; Maganuco, S.; Buffetaut, E.; Mendez, M. A. (2005). "New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its sizes and affinities". Journal of Vertebrate Paleontology (Submitted manuscript). 25 (4): 888–896. doi:10.1671/0272-4634(2005)025[0888:NIOTSO]2.0.CO;2.
  23. ^ http://www.awf.org/wildlife-conservation/ostrich
  24. ^ Conservation International (Content Partner); Mark McGinley (Topic Editor). 2008. "Biological diversity in the Caribbean Islands." In: Encyclopedia of Earth. Eds. Cutler J. Cleveland (Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment). [First published in the Encyclopedia of Earth May 3, 2007; Last revised August 22, 2008; Retrieved November 9, 2009]. <http://www.eoearth.org/article/Biological_diversity_in_the_Caribbean_Islands>
  25. ^ a b Borenstein, Seth (July 31, 2014). "Study traces dinosaur evolution into early birds". AP News. Retrieved August 3, 2014.
  26. ^ a b Zoe Gough (31 July 2014). "Dinosaurs 'shrank' regularly to become birds". BBC.
  27. ^ a b c d e Hutchinson, J.R. (March–April 2006). "The evolution of locomotion in archosaurs". Comptes Rendus Palevol. 5 (3–4): 519–530. doi:10.1016/j.crpv.2005.09.002. Archived from the original on 2008-12-01.
  28. ^ a b Newman, BH (1970). "Stance and gait in the flesh-eating Tyrannosaurus". Biological Journal of the Linnean Society. 2 (2): 119–123. doi:10.1111/j.1095-8312.1970.tb01707.x.
  29. ^ K. Padian, P.E. Olsen, (1989). "Ratite footprints and the stance and gait of Mesozoic theropods." Pp. 231–241 in: D.D. Gillette, M.G. Lockley (Eds.), Dinosaur Tracks and Traces, Cambridge University Press, Cambridge.
  30. ^ Paul, G.S. (1998). "Limb design, function and running performance in ostrich-mimics and tyrannosaurs". Gaia. 15: 257–270.
  31. ^ Farlow, J.O.; Gatesy, S.M.; Holtz, Jr.; Hutchinson, J.R.; Robinson, J.M. (2000). "Theropod locomotion". Am. Zool. 40 (4): 640–663. doi:10.1093/icb/40.4.640.
  32. ^ "Abstract", in Chure (2001). Pg. 19.
  33. ^ Dong, Z (1984). "A new theropod dinosaur from the Middle Jurassic of Sichuan Basin". Vertebrata PalAsiatica. 22 (3): 213–218.
  34. ^ a b c d e f Rauhut, O.W. (2003). The Interrelationships and Evolution of Basal Theropod Dinosaurs. Blackwell Publishing, 213 pp. ISBN 0-901702-79-X
  35. ^ See Origin of birds/Digit homology.
  36. ^ Some genera within Avetheropoda, however, had four digits, see University of Maryland department of geology home page, "Theropoda I" on Avetheropoda, 14 July 2006.
  37. ^ a b Carpenter, K. (2002). "Forelimb biomechanics of nonavian theropod dinosaurs in predation". Senckenbergiana Lethaea. 82 (1): 59–76. doi:10.1007/BF03043773.
  38. ^ a b Senter, P.; Robins, J.H. (July 2005). "Range of motion in the forelimb of the theropod dinosaur Acrocanthosaurus atokensis, and implications for predatory behaviour". Journal of Zoology, London. 266 (3): 307–318. doi:10.1017/S0952836905006989.
  39. ^ Molnar, R. E., 2001, Theropod paleopathology: a literature survey: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 337-363.
  40. ^ a b Xing, L D; Lockley, M G; Zhang, J P; et al. (2013). "A new Early Cretaceous dinosaur track assemblage and the first definite non-avian theropod swim trackway from China". Chin Sci Bull. 58 (19): 2370–2378. doi:10.1007/s11434-013-5802-6.
  41. ^ Nesbitt, S. J.; Smith, N. D.; Irmis, R. B.; Turner, A. H.; Downs, A. & Norell, M. A. (11 December 2009). "A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs". Science. 326 (5959): 1530–1533. doi:10.1126/science.1180350. PMID 20007898..
  42. ^ Rowe, T., and Gauthier, J., (1990). "Ceratosauria." Pp. 151–168 in Weishampel, D. B., Dodson, P., and Osmólska, H. (eds.), The Dinosauria, University of California Press, Berkeley, Los Angeles, Oxford.
  43. ^ Mortimer, M. (2001). "Rauhut's Thesis", Dinosaur Mailing List Archives, 4 Jul 2001.
  44. ^ Carrano, M. T.; Sampson, S. D.; Forster, C. A. (2002). "The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar". Journal of Vertebrate Paleontology. 22 (3): 510–534. doi:10.1671/0272-4634(2002)022[0510:TOOMKA]2.0.CO;2.
  45. ^ Ostrom, J.H. (1969). "Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana". Peabody Museum Natural History Bulletin. 30: 1–165.
  46. ^ Paul, G.S. (1988). Predatory Dinosaurs of the World. New York: Simon and Schuster Co. (ISBN 0-671-61946-2)
  47. ^ Dingus, L. and Rowe, T. (1998). The Mistaken Extinction: Dinosaur Evolution and the Origin of Birds. Freeman.
  48. ^ Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press. 472 pp. (ISBN 0-8018-6763-0)
  49. ^ Lee, MichaelS.Y.; Cau, Andrea; Naish, Darren; Dyke, Gareth J. (1 August 2014). "Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds". Science. 345 (6196): 562–566. doi:10.1126/science.1252243. PMID 25082702.
  50. ^ Marsh, O.C. (1881). "Principal characters of American Jurassic dinosaurs. Part V.". The American Journal of Science and Arts. 3. 21 (125): 417–423. doi:10.2475/ajs.s3-21.125.417.
  51. ^ Matthew, W. D.; Brown, B. (1922). "The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta". Bulletin of the American Museum of Natural History. 46: 367–385.
  52. ^ Anderson, Ted R. (2006). Biology of the Ubiquitous House Sparrow: from Genes to Populations. Oxford: Oxford University Press. ISBN 0-19-530411-X.
  53. ^ Holtz, Thomas R. Jr. (2011) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
  54. ^ Hendrickx, C.; Hartman, S.A.; Mateus, O. (2015). "An Overview of Non- Avian Theropod Discoveries and Classification". PalArch's Journal of Vertebrate Palaeontology. 12 (1): 1–73.
  55. ^ Baron, M.G.; Norman, D.B.; Barrett, P.M. (2017). "A new hypothesis of dinosaur relationships and early dinosaur evolution" (PDF). Nature. 543 (7646): 501–506. doi:10.1038/nature21700. PMID 28332513.
  56. ^ https://www.cam.ac.uk/research/news/new-study-shakes-the-roots-of-the-dinosaur-family-tree
Allosauroidea

Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among the apex predators that were active during the Middle Jurassic to Late Cretaceous periods. Of the fourteen allosauroid taxa, five are known for specimens with relatively complete skulls; the taxa are Allosaurus, Sinraptor, Yangchuanosaurus, Carchardontosaurus, and Acrocanthosaurus. The most famous and best understood allosauroid is the North American genus Allosaurus.

The oldest-known allosauroid, Shidaisaurus jinae, appeared in the early Middle Jurassic (probably Bajocian stage) of China. The last known definitive surviving members of the group died out around 93 million years ago in Asia (Shaochilong) and South America (Mapusaurus), though the megaraptorans may belong to the group as well. Additional, but highly fragmentary, remains probably belonging to carcharodontosaurids have been found from the Late Maastrichtian (70-66 Ma ago) in Brazil. An alternative interpretation is to attribute the remains to abelisaurids, which share the distinct pattern of curved wrinkled enamel found in the Brazilian remains with the carcharodontosaurids. This similarity between abelisaurids and carcharodontosaurids means that a definitive match between the Brazilian fossil and carcharodontosaurids cannot be made.Allosauroids had long, narrow skulls, large orbits, three-fingered hands, and usually had "horns" or ornamental crests on their heads. Although allosauroids vary in size, the group maintains a similar center of mass and hip position on their bodies. Allosauroids also exhibit reptilian-style immune systems, secreting fibrin at injured sites to prevent infections from spreading through the bloodstream. This characteristic has been observed by examining injuries and infections on allosauroid bones. It is possible that allosauroids were social animals, as many remains of allosauroids have been found in close proximity to each other. Allosauroids were likely active predators, and from studying endocasts, probably best responded to odors and loud low-frequency noises.

Averostra

Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.

Avetheropoda

Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.

Barremian

The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 125.0 ± 1.0 Ma). It is a subdivision of the Early Cretaceous epoch (or Lower Cretaceous series). It is preceded by the Hauterivian and followed by the Aptian stage.

Cenomanian

The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding age. Both age and stage bear the same name.

As a unit of geologic time measure, the Cenomanian age spans the time between 100.5 ± 0.9 Ma and 93.9 ± 0.8 Ma (million years ago). In the geologic timescale it is preceded by the Albian and is followed by the Turonian. The Upper Cenomanian starts approximately at 95 M.a.

The Cenomanian is coeval with the Woodbinian of the regional timescale of the Gulf of Mexico and the early part of the Eaglefordian of the regional timescale of the East Coast of the United States.

At the end of the Cenomanian an anoxic event took place, called the Cenomanian-Turonian boundary event or the "Bonarelli Event", that is associated with a minor extinction event for marine species.

Chuandongocoelurus

Chuandongocoelurus ( chwahn-DONG-ə-si-LEWR-əs) is a genus of carnivorous tetanuran theropod dinosaur from the Jurassic of China.

Coelurosauria

Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.

Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaur group alive today.Most feathered dinosaurs discovered so far have been coelurosaurs. Philip J. Currie considers it likely and probable that all coelurosaurs were feathered. In the past, Coelurosauria was used to refer to all small theropods, but this classification has since been abolished.

Como Bluff

Como Bluff is a long ridge extending east-west, located between the towns of Rock River and Medicine Bow, Wyoming. The ridge is an anticline, formed as a result of compressional geological folding. Three geological formations, the Sundance, the Morrison, and the Cloverly Formations, containing fossil remains from the Late Jurassic of the Mesozoic Era are exposed. Nineteenth century paleontologists discovered many well-preserved specimens of dinosaurs, as well as mammals, turtles, crocodilians, and fish from the Morrison Formation. Because of this, Como Bluff is considered to be one of the major sites for the early discovery of dinosaur remains. Among the species discovered is the only known specimen of Coelurus. Significant discoveries were made in 22 different areas scattered along the entire length of the ridge. It is included on the National Register of Historic Places as well as the National Natural Landmark list.

Dinosauriformes

Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.

Evolution of dinosaurs

This article gives an outline and examples of dinosaur evolution. For a detailed list of interrelationships see Dinosaur classification.

Dinosaurs evolved within a single lineage of archosaurs 243-233 Ma (million years ago) from the Anisian to the Carnian ages, the latter part of the middle Triassic. Dinosauria is a well-supported clade, present in 98% of bootstraps. It is diagnosed by many features including loss of the postfrontal on the skull and an elongate deltopectoral crest on the humerus.In March 2017, scientists reported a new way of classifying the dinosaur family tree, based on newer and more evidence than available earlier. According to the new classification, the original dinosaurs, arising 200 million years ago, were small, two-footed omnivorous animals with large grasping hands. Descendants (for the non-avian dinosaurs) lasted until 66 million years ago.

Garden Park, Colorado

Garden Park is a paleontological site in Fremont County, Colorado, known for its Jurassic dinosaurs and the role the specimens played in the infamous Bone Wars of the late 19th century. Located 10 km (6.2 mi) north of Cañon City, the name originates from the area providing vegetables to the miners at nearby Cripple Creek in the 19th century. Garden Park proper is a triangular valley surrounded by cliffs on the southeast and southwest and by mountains to the north; however, the name is also refers to the dinosaur sites on top and along the cliffs. The dinosaur sites now form the Garden Park Paleontological Resource Area, which is overseen by the Bureau of Land Management.

Majungasaurus

Majungasaurus (; "Mahajanga lizard") is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period. The genus contains a single species, Majungasaurus crenatissimus. This dinosaur was briefly called Majungatholus, a name which is now considered a junior synonym of Majungasaurus.

Like other abelisaurids, Majungasaurus was a bipedal predator with a short snout. Although the forelimbs are not completely known, they were very short, while the hind limbs were longer and very stocky. It can be distinguished from other abelisaurids by its wider skull, the very rough texture and thickened bone on the top of its snout, and the single rounded horn on the roof of its skull, which was originally mistaken for the dome of a pachycephalosaur. It also had more teeth in both upper and lower jaws than most abelisaurids.

Known from several well-preserved skulls and abundant skeletal material, Majungasaurus has recently become one of the best-studied theropod dinosaurs from the Southern Hemisphere. It appears to be most closely related to abelisaurids from India rather than South America or continental Africa, a fact that has important biogeographical implications. Majungasaurus was the apex predator in its ecosystem, mainly preying on sauropods like Rapetosaurus, and is also one of the few dinosaurs for which there is direct evidence of cannibalism.

Megalosauroidea

Megalosauroidea (meaning 'great/big lizard forms') is a superfamily (or clade) of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as Megalosaurus bucklandii and all taxa sharing a more recent common ancestor with it than with Allosaurus fragilis or Passer domesticus. Members of the group include Spinosaurus, Megalosaurus, and Torvosaurus.

Neotheropoda

Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.

Orionides

Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.

Sarcosaurus

Sarcosaurus (meaning "flesh lizard") is a genus of theropod dinosaur, roughly 3.5 metres (11 ft) long. It lived during the Sinemurian stage of the Early Jurassic, about 194 million years ago. Sarcosaurus is one of the earliest known Jurassic theropods, and one of only a handful of theropod genera from this time period.

Saurischia

Saurischia ( saw-RIS-kee-ə, meaning "reptile-hipped" from the Greek sauros (σαῦρος) meaning 'lizard' and ischion (ἴσχιον) meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia). ‘Saurischia’ translates to lizard-hipped.

In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure, though today most paleontologists classify Saurischia as an unranked clade rather than an order.

Tetanurae

Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, maniraptorans, and birds. Tetanurans are defined as all theropods more closely related to modern birds than to Ceratosaurus and contain the majority of predatory dinosaur diversity. Tetanurae likely diverged from its sister group, Ceratosauria, during the late Triassic. Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.The group was named by Jacques Gauthier in 1986 and originally had two main subgroups: Carnosauria and Coelurosauria, the clade containing birds and related dinosaurs such as compsognathids, tyrannosaurids, ornithomimosaurs, and maniraptorans. The original Carnosauria was a polyphyletic group including any large carnivorous theropod. Many of Gauthier's carnosaurs, such as tyrannosaurids, have since been re-classified as coelurosaurs or primitive tetanurans. Carnosauria has been reclassified as a group containing allosaurids that split from the Coelurosauria at the Neotetanurae/Avetheropoda node. Members of Spinosauroidea are believed to represent basal tetanurans.Tetanuran evolution was characterized by parallel diversification of multiple lineages, repeatedly attaining large body size and similar locomotor morphology. Cryolophosaurus has been claimed as the first true member of the group, but subsequent studies have disagreed on whether it is a dilophosaurid or tetanuran. Arcucci and Coria (2003) classified Zupaysaurus as an early tetanuran, but it was later placed as a sister taxon to the clade containing dilophosaurids, ceratosaurs, and tetanurans.Shared tetanuran features include a ribcage indicating a sophisticated air-sac-ventilated lung system similar to that in modern birds. This character would have been accompanied by an advanced circulatory system. Other tetanuran characterizing features include the absence of the fourth digit of the hand, placement of the maxillary teeth anterior to the orbit, a strap-like scapula, maxillary fenestrae, and stiffened tails. During the Late Jurassic and Early Cretaceous, large spinosaurids and allosaurs flourished but possibly died out in the northern hemisphere before the end of the Cretaceous, and were replaced as apex predators by tyrannosauroid coelurosaurs. At least in South America, carcharodontosaurid allosaurs persisted until the end of the Mesozoic Era, and died out at the same time the non-avian coelurosaurs.

Languages

This page is based on a Wikipedia article written by authors (here).
Text is available under the CC BY-SA 3.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.