Theropoda

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Not to be confused with Pteropoda, marine gastropods.
Theropods
Temporal range:
Late Triassic – Present, 231.4–0 Ma
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Theropoda 2.png
Theropod morphological and ecological diversity
Scientific classificatione
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Eusaurischia
Clade: Theropoda
Marsh, 1881
Subgroups[1]
†Daemonosaurus?
†Eodromaeus?
†Erythrovenator
†Guaibasaurus?
†Nhandumirim?
†Tawa
†Velocipes?
Neotheropoda
Theropoda (/θɪəˈrɒpədə/[2] from Greek θηρίον 'wild beast' and πούς, ποδός 'foot'),
whose members are known as theropods, is a dinosaur clade that is characterized by
hollow bones and three-toed limbs. Theropods are generally classed as a group of
saurischian dinosaurs. They 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)[3] 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.

Contents
1 Biology
1.1 Diet and teeth
1.2 Integument (skin, scales and feathers)
1.3 Size
1.4 Growth rates
1.5 Stance and gait
1.6 Nervous system and senses
1.7 Forelimb morphology
1.8 Forelimb movement
1.9 Paleopathology
1.10 Swimming
2 Evolutionary history
3 Classification
3.1 History of classification
3.2 Major groups
3.3 Relationships
4 See also
5 References
6 External links
Biology
Diet and teeth

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.[4] 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 maniraptorans 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.[5][6]

Diet is largely deduced by the tooth morphology,[7] 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,[8]
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,[7] 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.[9] The folds helped the teeth stay in place
longer, especially as theropods evolved into larger sizes and had more force in
their bite.[10][11]

Integument (skin, scales and feathers)

Fossil of an Anchiornis, showing large preserved feather imprints

Mesozoic theropods were also very diverse in terms of skin texture and covering.
Feathers or feather-like structures (filaments) 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.[12]

The coelurosaur lineages most distant from birds had feathers that were relatively
short and composed of simple, possibly branching filaments.[13] Simple filaments
are also seen in therizinosaurs, which also possessed large, stiffened "quill"-like
feathers. More fully feathered theropods, such as dromaeosaurids, 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,[14] and
Juravenator may have been predominantly scaly with some simple filaments
interspersed.[15] 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.[16]

Size
Main article: Dinosaur size
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.[17] 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 more massive than Spinosaurus.[18] Specimens such as
Sue and Scotty are both estimated to be the heaviest theropods known to science.
There is still no clear explanation for exactly why these animals grew so heavy and
bulky compared to 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 90 and 130 kg (200 - 290 lb).[19]

Size comparison of selected giant theropod dinosaurs

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.[16]
When modern birds are included, the bee hummingbird Mellisuga helenae is smallest
at 1.9 g and 5.5 cm (2.2 in) long.[20]

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.
[21][22]

Growth rates
In order to estimate the growth rates of theropods, scientists need to calculate
both age and body mass. Both of these measures can only be calculated through
fossilized bone and tissue, so regression analysis and extant animal growth rates
as proxies are used to make predictions. Fossilized bones exhibit growth rings that
appear as a result of growth or seasonal changes, which can be used to approximate
age at the time of death.[23] However, the amount of rings in a skeleton can vary
from bone to bone, and old rings can also be lost at advanced age, so scientists
need to properly control these two possibly confounding variables.

Body mass is harder to determine as bone mass only represents a small proportion of
the total body mass of animals. One method is to measure the circumference of the
femur, which in non-avian theropod dinosaurs has been shown to be a relatively
proportional to quadrupedal mammals,[24] and use this measurement as a function of
body weight, as the proportions of long bones like the femur grow proportionately
with body mass.[24] The method of using extant animal bone proportion to body mass
ratios to make predictions about extinct animals is known as the extant-scaling
(ES) approach.[25] A second method, known as the volumetric-density (VD) approach,
uses full scale models of skeletons to make inferences about potential mass.[25]
The ES approach is better for wide range studies including many specimens and
doesn't require as much of a complete skeleton as the VD approach, but the VD
approach allows scientists to better answer more physiological questions about the
animal, such as locomotion and center of gravity.[25]

The current consensus is that non-avian theropods didn't exhibit a group wide
growth rate, but instead had varied rates depending on their size. However, all
non-avian theropods had faster growth rates than extant reptiles, even when modern
reptiles are scaled up to the large size of some non-avian theropods. As body mass
increases, the relative growth rate also increases. This trend may be due to the
need to reach the size required for reproductive maturity.[26] For example, one of
the smallest known theropods was Microraptor zhaoianus, which had a body mass of
200 grams, grew at a rate of approximately .33 grams per day.[27] A comparable
reptile of the same size grows at half of this rate.[27] The growth rates of medium
sized non-avian theropods (100–1000 kg) approximated those of precocial birds,
which are much slower than altricial birds. Large theropods (1500–3500 kg) grew
even faster, similar to rates displayed by eutherian mammals.[27] The largest non-
avian theropods, like Tyrannosaurus rex had similar growth dynamics to the largest
living land animal today, the African elephant, which is characterized by a rapid
period of growth until maturity, subsequently followed by slowing growth in
adulthood.[28]

Stance and gait

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.[29] 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.[29]

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.[29]
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.[29][30] 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,[30] while
others suggest that the knee was normally strongly flexed in all theropods while
walking, even giants like the tyrannosaurids.[31][32] It is likely that a wide
range of body postures, stances, and gaits existed in the many extinct theropod
groups.[29][33]

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."[34]
Forelimb morphology

Mummified enantiornithean wing (of an unknown genus) from Cenomanian amber from
Myanmar
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.[35] However, this is no longer thought to be likely.[36]

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), 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.[37]

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.[14] 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

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.[38] 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.[39] 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.[38]

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. 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 dromaeosaurids, the hand
itself had lost most flexibility, with highly inflexible fingers. Dromaeosaurids
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.[39]

Paleopathology
Main article: Theropod 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.[40]

Swimming
The trackway of a swimming theropod, the first in China of the ichnogenus named
Characichnos, was discovered at the Feitianshan Formation in Sichuan.[41] 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.[41]

Evolutionary history
Full skeleton of an early carnivorous dinosaur, displayed in a glass case in a
museum
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.[42]

The earliest and most primitive unambiguous theropods (or alternatively,

"Eutheropoda"—'True Theropods') are the Coelophysoidea. The coelophysoids were a
group of widely distributed, lightly built and potentially gregarious animals. They
included small hunters like Coelophysis and Camposaurus. 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,[43]
they may have been ancestral to all other theropods (which would make them a
paraphyletic group).[44][45]

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[46][47]), 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.[48] While
the roots of these various groups are found in the Middle Jurassic, they only
became abundant during the Early Cretaceous. A few palaeontologists, 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.[49]

On July 31, 2014, scientists reported details of the evolution of birds from other
theropod dinosaurs.[21][22][50] 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, who coined the name Theropoda. Photo c. 1870
O. C. Marsh coined the name Theropoda (meaning "beast feet") in 1881.[51] 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").[36]

By the early 20th century, some palaeontologists, 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).[36] In W. D. Matthew and Barnum Brown's
1922 description of the first known dromaeosaurid (Dromaeosaurus albertensis[52]),
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.[36]

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 palaeontology, 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.[36]

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.[36]

Major groups

Ceratosaurus, a ceratosaurid

Irritator, a spinosaurid

Mapusaurus, a carcharodontosaurid

Microraptor, a dromaeosaurid

Passer domesticus, an avian, and the world's most widespread extant wild theropod.
[53]
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.[1] A more detailed version can be found at dinosaur
classification. The dagger (†) is used to signify groups with no living members.

†Herrerasauria (early bipedal carnivores)

†Coelophysoidea (small, early theropods; includes Coelophysis and close relatives)
†Dilophosauridae (early crested and carnivorous theropods)
†Ceratosauria (generally elaborately horned, the dominant southern carnivores of
the Cretaceous)
Tetanurae ("stiff tails"; includes most theropods)
†Megalosauroidea (early group of large carnivores including the semi-aquatic
spinosaurids)
†Carnosauria (Allosaurus and close relatives, like Carcharodontosaurus)
Coelurosauria (feathered theropods, with a range of body sizes and niches)
†Compsognathidae (early coelurosaurs with reduced forelimbs)
†Tyrannosauridae (Tyrannosaurus and close relatives; had reduced forelimbs)
†Ornithomimosauria ("ostrich-mimics"; mostly toothless; carnivores to possible
herbivores)
Maniraptora ("hand snatchers"; had long, slender arms and fingers)
†Alvarezsauroidea (small insectivores with reduced forelimbs each bearing one
enlarged claw)
†Therizinosauria (bipedal herbivores with large hand claws and small heads)
†Scansoriopterygidae (small, arboreal maniraptors with long third fingers)
†Oviraptorosauria (mostly toothless; their diet and lifestyle are uncertain)
†Archaeopterygidae (small, winged protobirds)
†Dromaeosauridae (small to medium-sized theropods)
†Troodontidae (small, gracile theropods)
Avialae (birds and extinct relatives)
†Omnivoropterygidae (large, early short-tailed avialans)
†Confuciusornithidae (small toothless birds)
†Enantiornithes (primitive tree-dwelling, flying birds)
Euornithes (advanced flying birds)
†Yanornithiformes (toothed Cretaceous Chinese birds)
†Hesperornithes (specialized aquatic diving birds)
Aves (modern, beaked birds and their extinct relatives)