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{{Short description|Archosaurian reptiles that dominated the Mesozoic Era}}
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{{Taxobox | color = pink
{{Distinguish|Dinosaurus}}
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{{Use mdy dates|date=January 2022}}
{{Automatic taxobox
| name = Dinosaurs | name = Dinosaurs
| fossil_range = ] – ] (except avian) | fossil_range = {{fossilrange|237|0|]–], 233.23 0 Mya | |earliest=243 |PS=(range includes ]s) }} (possible Middle Triassic record)
| image = Saurier2.jpg | image = {{Multiple image
| perrow = 2/2/2
| image_width = 260px
| width = 145
| image_caption = Replica of '']'' at the ].
| caption_align = center
| regnum = ]ia
| image1 = Frenguellisaurus ischigualastensis DSC 6185.jpg
| phylum = ]
| caption1 = '']''<br />(a carnivorous basal dinosaur)
| classis = ]
| image2 = Triceratops Specimen at the Houston Museum of Natural Science.JPG
| subclassis = ]a
| caption2 = '']''<br />(a ]n)
| infraclassis = ]
| image3 = Stegosaurus ungulatus.jpg
{{Taxobox_norank_entry | taxon = ]}}
| caption3 = '']''<br />(a ])
{{Taxobox_norank_entry | taxon = ]}}
| image4 = Louisae.jpg
| superordo = '''Dinosauria''' ]
| caption4 = '']''<br />(a ])
| superordo_authority = ], 1842
| image5 = Hadrosauridae - Edmontosaurus annectens.JPG
| subdivision_ranks = Orders & Suborders
| caption5 = '']''<br />(a ] ])
| subdivision = <div>
| image6 = MicroraptorGui-PaleozoologicalMuseumOfChina-May23-08.jpg
* ''']'''
| caption6 = '']''<br />(a ] ])
** ]
| border = infobox
** ]
* ''']'''
** ]
** ]
** ]
</div>
}} }}
| display_parents = 7
'''Dinosaurs''' were ] ]s that dominated ] ]s for over 160 million years, first appearing approximately 230 ]. At the end of the ] ], 65 million years ago, dinosaurs suffered a catastrophic ], which ended their dominance on land. ] consider modern birds to be the direct descendants of theropod dinosaurs.
| taxon = Dinosauria
| authority = ], 1842
| subdivision_ranks = Major groups
| subdivision = *{{extinct}}]
*{{extinct}}]
*]
**Various extinct groups
**] (birds)
{{Collapse top|title=Dinosaurs and possible dinosaurs of uncertain affinity|left=yes|padding=0|border=0|border2=0|bg=clear|bg2=clear}}
*{{extinct}}'']''?
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*{{extinct}}'']''?
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*{{extinct}}]<ref>Matthew G. Baron; Megan E. Williams (2018). "A re-evaluation of the enigmatic dinosauriform Caseosaurus crosbyensis from the Late Triassic of Texas, USA and its implications for early dinosaur evolution". Acta Palaeontologica Polonica. 63. {{doi|10.4202/app.00372.2017}}.</ref><ref>Andrea Cau (2018). "The assembly of the avian body plan: a 160-million-year long process" (PDF). Bollettino della Società Paleontologica Italiana. 57 (1): 1–25. {{doi|10.4435/BSPI.2018.01}}.</ref>
*{{extinct}}]? <small>(]?)</small><ref>{{Cite journal|last1=Ferigolo|first1=Jorge|last2=Langer |first2=Max C. |date=2007|title=A Late Triassic dinosauriform from south Brazil and the origin of the ornithischian predentary bone |journal=Historical Biology |volume=19|issue=1|pages=23–33 |doi=10.1080/08912960600845767 |bibcode=2007HBio...19...23F |s2cid=85819339|issn=0891-2963}}</ref><ref>{{Cite journal |last1=Langer |first1=Max C. |last2=Ferigolo |first2=Jorge |date=2013 |title=The Late Triassic dinosauromorph ''Sacisaurus agudoensis'' (Caturrita Formation; Rio Grande do Sul, Brazil): anatomy and affinities |url=https://sp.lyellcollection.org/content/379/1/353 |journal=Geological Society, London, Special Publications |language=en |volume=379 |issue=1 |pages=353–392 |doi=10.1144/SP379.16 |bibcode=2013GSLSP.379..353L |s2cid=131414332|issn=0305-8719}}</ref><ref name="cabreira2016">{{cite journal |last1=Cabreira |first1=S.F. |last2=Kellner |first2=A.W.A.|last3=Dias-da-Silva|first3=S.|last4=da Silva |first4=L.R. |last5=Bronzati |first5=M. |last6=de Almeida Marsola |first6=J.C. |last7=Müller|first7=R.T.|last8=de Souza Bittencourt |first8=J. |last9=Batista |first9=B.J. |last10=Raugust |first10=T. |last11=Carrilho |first11=R.|date=2016|title=A Unique Late Triassic Dinosauromorph Assemblage Reveals Dinosaur Ancestral Anatomy and Diet |journal=Current Biology|volume=26|issue=22|pages=3090–3095 |doi=10.1016/j.cub.2016.09.040 |pmid=27839975 |doi-access=free |first12=A. |last12=Brodt |first13=M.C.|last13=Langer|bibcode=2016CBio...26.3090C | issn = 0960-9822}}</ref><ref>{{Cite journal |last1=Müller|first1=Rodrigo Temp |last2=Garcia |first2=Maurício Silva |date=August 26, 2020|title=A paraphyletic 'Silesauridae' as an alternative hypothesis for the initial radiation of ornithischian dinosaurs |journal=Biology Letters |volume=16 |issue=8 |pages=20200417 |doi=10.1098/rsbl.2020.0417 |pmid=32842895|pmc=7480155|doi-access=free}}</ref>
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]s are avian dinosaurs, and in ] ] their over 11,000 extant species are included in the group Dinosauria.]]
'''Dinosaurs''' are<!-- Please use "are", as the scientific consensus is that birds are also dinosaurs themselves. --> a diverse group of ]s{{refn|group=note|name=cold-blooded|Dinosaurs (including birds) are members of the ] ]. Their biology does not precisely correspond to the antiquated ] Reptilia of ], consisting of ] ]s without fur or feathers. As Linnean taxonomy was formulated for modern animals prior to the study of evolution and paleontology, it fails to account for extinct animals with intermediate traits between traditional classes.}} of the ] '''Dinosauria'''. They first appeared during the ] ], between 243 and 233.23&nbsp;] (mya), although the exact origin and timing of the ] is a subject of active research. They became the dominant terrestrial ]s after the ] 201.3&nbsp;mya and their dominance continued throughout the ] and ] periods. The ] record shows that ]s are ]s, ] from earlier ] during the ] ], and are the only dinosaur lineage known to have survived the ] approximately 66&nbsp;mya. Dinosaurs can therefore be divided into '''avian dinosaurs'''—birds—and the extinct '''non-avian dinosaurs''', which are all dinosaurs other than birds.


Dinosaurs are varied from ], ] and ] standpoints. Birds, at over 11,000 living ], are among the most diverse groups of vertebrates. Using fossil evidence, ] have identified over 900 distinct ] and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both ] species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized as dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and ]. Most ], however, has indicated that dinosaurs were active animals with elevated ]s and numerous adaptations for social interaction. Some were ], others ]. Evidence suggests that all dinosaurs were ], and that ]-building was a trait shared by many dinosaurs, both avian and non-avian.
Since the first dinosaur was recognized in the 19th century, mounted, fossilized dinosaur skeletons have become major attractions at ]s around the world. Dinosaurs have become a part of world culture and remain consistently popular, especially among children. They have been featured in best-selling books and films such as '']'', and new discoveries are regularly covered by the ].


While dinosaurs were ancestrally ], many ] groups included ] species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed ] modifications such as ] and ]. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest ] dinosaurs are estimated to have reached lengths of {{convert|39.7|m|ft|abbr=off|sp=us}} and heights of {{convert|18|m|ft|abbr=on|sp=us}} and were the largest land animals of all time. The misconception that non-avian dinosaurs were uniformly gigantic is based in part on ], as large, sturdy ]s are more likely to last until they are fossilized. Many dinosaurs were quite small, some measuring about {{convert|50|cm|in|abbr=off|sp=us}} in length.
The term ''dinosaur'' is sometimes used informally to describe other prehistoric reptiles, such as the ] '']'', the winged ]s and the aquatic ]s, ]s and ]s, although technically none of these were dinosaurs.


The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" (meaning "terrible lizard") being coined by Sir ] in 1842 to refer to these "great fossil lizards".<ref name=":2">{{Cite web |date=2017-04-28 |title=The 'birth' of dinosaurs |url=https://morethanadodo.com/2017/04/28/the-birth-of-dinosaurs/ |access-date=2023-03-15 |website=More Than A Dodo}}</ref><ref name=":3">{{Cite news |date=October 16, 2015 |title=The Birth of Dinosaurs: Richard Owen and Dinosauria |url=https://blog.biodiversitylibrary.org/2015/10/the-birth-of-dinosaurs-richard-owen-and-dinosauria.html |access-date=March 15, 2023 |website=Biodiversity Heritage Library }}</ref><ref name=":4">{{Cite book |last1=Brett-Surman |first1=M. K. |url=https://books.google.com/books?id=pX_l24sDARwC&dq=Hugh+Torrens+Dinosauria+Owen&pg=PA25 |title=The Complete Dinosaur |last2=Holtz |first2=Thomas R. |last3=Farlow |first3=James O. |date=June 27, 2012 |publisher=Indiana University Press |isbn=978-0-253-00849-7 |page=25 }}</ref> Since then, mounted fossil dinosaur skeletons have been major attractions at museums worldwide, and dinosaurs have become ]. The large sizes of some dinosaurs, as well as their seemingly monstrous and fantastic nature, have ensured their regular appearance in best-selling books and films, such as the '']'' franchise. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.
==What is a dinosaur?==
===Definition===
]'' ] at the ] ].]]
The ] '''Dinosauria''' was formally named by the ] ] ] in 1842 as "a distinct tribe or suborder of Saurian reptiles".<ref>* Owen, Richard. 1842. "Report on British Fossil Reptiles." Part II. Report of the British Association for the Advancement of Science, Plymouth, England.</ref> The term is derived from the ] words δεινός (''deinos'' meaning "terrible", "fearsome" or "formidable") and σαύρα (''saura'' meaning "lizard" or "reptile"). Owen chose it to express his awe at the size and majesty of the extinct animals, not out of fear or trepidation at their size and often-formidable arsenal of teeth and claws.


==Definition==
Dinosaurs were an extremely varied group of animals; according to a 2006 study, 527 dinosaur genera have been identified with certainty so far, and 1,844 genera are believed to have existed.<ref>Fountain, Henry. "Many more dinosaurs still to be found." ''New York Times'': 12 Sept. 2006. </ref><ref>Wang, S.C., and Dobson, P. (2006). Estimating the Diversity of Dinosaurs. Proceedings of the National Academy of Sciences USA 103:37, pp. 13601-13605. </ref> Some were ], others ]. Some dinosaurs were ]s, some were ]s and others, such as '']'' and '']'', could walk just as easily on two or four legs. Regardless of body type, nearly all known dinosaurs were well-adapted for a predominantly terrestrial, rather than aquatic or aerial, habitat.
Under ], dinosaurs are usually defined as the group consisting of the ] (MRCA) of '']'' and ] (Neornithes), and all its descendants.<ref name=MJB04dino/> It has also been suggested that Dinosauria be defined with respect to the MRCA of '']'' and '']'', because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria.<ref name=olshevsky2000/> Both definitions cover the same known genera: Dinosauria = ] + ]. This includes major groups such as ]ns (armored herbivorous quadrupeds), ]ns (plated herbivorous quadrupeds), ]ns (bipedal or quadrupedal herbivores with ]s), ]ns (bipedal herbivores with thick skulls), ]s (bipedal or quadrupedal herbivores including "]"), ]s (mostly bipedal carnivores and birds), and ] (mostly large herbivorous quadrupeds with long necks and tails).<ref name="Letal05"/>


Birds are the sole surviving dinosaurs. In traditional ], birds were considered a separate ] that had evolved from dinosaurs, a distinct ]. However, most contemporary paleontologists reject the traditional style of classification based on anatomical similarity, in favor of ] taxonomy based on deduced ancestry, in which each group is defined as all descendants of a given founding genus.<ref>{{cite web |url=https://evolution.berkeley.edu/evolibrary/article/evo_10 |url-status=live |title=Using the tree for classification |website=Understanding Evolution |publisher=] |location=] |archive-url=https://web.archive.org/web/20190831003846/https://evolution.berkeley.edu/evolibrary/article/evo_10 |archive-date=August 31, 2019 |access-date=October 14, 2019}}</ref> Birds belong to the dinosaur subgroup ], which are ], which are theropods, which are saurischians.<ref name=KP04/>
'''Dinosaur ]'''


Research by Matthew G. Baron, ], and Paul M. Barrett in 2017 suggested a radical revision of dinosaurian systematics. Phylogenetic analysis by Baron ''et al.'' recovered the Ornithischia as being closer to the Theropoda than the Sauropodomorpha, as opposed to the traditional union of theropods with sauropodomorphs. This would cause sauropods and kin to fall outside traditional dinosaurs, so they re-defined Dinosauria as the last common ancestor of ''Triceratops horridus'', '']'' and '']'', and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. They also resurrected the clade ] to refer to the group containing Ornithischia and Theropoda.<ref name="NYT-20170322">{{cite news |last=Wade |first=Nicholas |author-link=Nicholas Wade |title=Shaking Up the Dinosaur Family Tree |date=March 22, 2017 |url=https://www.nytimes.com/2017/03/22/science/dinosaur-family-tree.html |url-status=live |url-access=registration |work=] |location=] |issn=0362-4331 |archive-url=https://web.archive.org/web/20180407234942/https://www.nytimes.com/2017/03/22/science/dinosaur-family-tree.html |archive-date=April 7, 2018 |access-date=October 30, 2019}} "A version of this article appears in print on March 28, 2017, on Page D6 of the New York edition with the headline: Shaking Up the Dinosaur Family Tree."</ref><ref name="NAT-20170322">{{cite journal |last1=Baron |first1=Matthew G. |last2=Norman |first2=David B. |author-link2=David B. Norman |last3=Barrett |first3=Paul M. |year=2017 |title=A new hypothesis of dinosaur relationships and early dinosaur evolution |journal=] |location=London |publisher=] |volume=543 |issue=7646 |pages=501–506 |bibcode=2017Natur.543..501B |doi=10.1038/nature21700 |issn=0028-0836 |pmid=28332513 |s2cid=205254710}}</ref>
All dinosaurs so far discovered share certain modifications to the ancestral ]ian skeleton. Although some later groups of dinosaurs featured further modified versions of these traits, they are considered typical across Dinosauria; the earliest dinosaurs had them and passed them on to all their descendants. Such common structures across a taxonomic group are called synapomorphies.


===General description===
Dinosaur synapomorphies include: reduced fourth and fifth digits on the ] ('hand'), reduced number of digits on the ] (foot) to three main toes, a sacrum (the region of the vertebral column to which the pelvis attaches, composed of three or more fused vertebrae) and an open or perforate ] (hip socket with a hole at its centre). Dinosaurs are unique among all ]s in having this perforate acetabulum.
]'' skeleton, ]]]


Using one of the above definitions, dinosaurs can be generally described as ]s with ].<ref name=DFG97/> Other prehistoric animals, including ]s, ]s, ]s, ], and '']'', while often popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs, being members of the clade ]. The other groups mentioned are, like dinosaurs and pterosaurs, members of ] (the reptile and bird clade), except ''Dimetrodon'' (which is a ]). None of them had the erect hind limb posture characteristic of true dinosaurs.<ref name=DL90/>
'''Other shared anatomical features'''


Dinosaurs were the dominant terrestrial vertebrates of the ] ], especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; ]s, for example, rarely exceeded the size of a domestic cat and were generally rodent-sized carnivores of small prey.<ref name=MM97/> Dinosaurs have always been recognized as an extremely varied group: over 900 non-avian dinosaur genera have been confidently identified (2018) with 1124 species (2016). Estimates put the total number of dinosaur genera preserved in the fossil record at 1850, nearly 75% still undiscovered,<ref name="Genera900">{{cite journal |last1=Tennant |first1=Jonathan P. |last2=Chiarenza |first2=Alfio Alessandro |last3=Baron |first3=Matthew |title=How has our knowledge of dinosaur diversity through geologic time changed through research history? |journal=PeerJ |date=19 February 2018 |volume=6 |pages=e4417 |doi=10.7717/peerj.4417|pmid=29479504 |pmc=5822849 |doi-access=free }}</ref><ref name="Howmany">{{cite journal |last1=Starrfelt |first1=Jostein |last2=Liow |first2=Lee Hsiang |title=How many dinosaur species were there? Fossil bias and true richness estimated using a Poisson sampling model |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |date=2016 |volume=371 |issue=1691 |pages=20150219 |doi=10.1098/rstb.2015.0219|pmid=26977060 |pmc=4810813 }}</ref><ref name="Wang&Dodson"/> and the number that ever existed (in or out of the fossil record) at 3,400.<ref name=russell1995/> A 2016 estimate put the number of dinosaur species living in the Mesozoic at 1,543–2,468,<ref>{{cite journal |last1=Starrfelt |first1=Jostein |last2=Liow |first2=Lee Hsiang |year=2016 |title=How many dinosaur species were there? Fossil bias and true richness estimated using a Poisson sampling model |journal=] |location=London |publisher=] |volume=371 |issue=1691 |page=20150219 |doi=10.1098/rstb.2015.0219 |issn=0962-8436 |pmc=4810813 |pmid=26977060}}</ref><ref>{{cite web |url=https://www.nationalgeographic.com/science/article/most-dinosaur-species-are-still-undiscovered/|url-status=live |url-access=registration |last=Black |first=Riley |title=Most Dinosaur Species Are Still Undiscovered |website=] |date=March 23, 2016 |archive-url=https://web.archive.org/web/20210306214843/https://www.nationalgeographic.com/science/article/most-dinosaur-species-are-still-undiscovered/ |archive-date=March 6, 2021 |access-date=June 6, 2021}}</ref> compared to the number of modern-day birds (avian dinosaurs) at 10,806 species.<ref>{{cite journal|last1=Gill|first1=F.|last2=Donsker|first2=D.|last3=Rasmussen|first3=P.|title=Welcome|year=2021|journal=IOC World Bird List 11.1.|url=https://www.worldbirdnames.org/new/}}</ref>
Scientists generally agree that a variety of other anatomical features were shared by most dinosaurs. These include forelimbs shorter and lighter than hind limbs, an unusual secondary palate that permitted dinosaurs to eat and breathe simultaneously, a relatively straight femur with medially-directed femoral head, two pairs of holes in the temporal region of the skull (i.e. a ] skull), rearward-pointing elbows in the front limbs and forward-pointing knees in the hind limbs.


Extinct dinosaurs, as well as modern birds, include genera that are herbivorous and others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some evolved into quadrupeds, and others, such as '']'' and ''Iguanodon'', could walk as easily on two or four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony armor. Although the best-known genera are remarkable for their large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global distribution by the ] epoch at latest.<ref name=MacLeod>{{cite journal |last1=MacLeod |first1=Norman |last2=Rawson |first2=Peter F. |last3=Forey |first3=Peter L. |last4=Banner |first4=FT |last5=Boudagher-Fadel |first5=MK |last6=Bown |first6=PR |last7=Burnett |first7=JA |last8=Chambers |first8=P |last9=Culver |first9=S |last10=Evans |first10=SE |last11=Jeffery |first11=C |last12=Kaminski |first12=MA |last13=Lord |first13=AR |last14=Milner |first14=AC |last15=Milner |first15=AR |last16=Morris |first16=N |last17=Owen |first17=E |last18=Rosen |first18=BR |last19=Smith |first19=AB |last20=Taylor |first20=PD |last21=Urquhart |first21=E |last22=Young |first22=JR |display-authors=3 |year=1997 |title=The Cretaceous–Tertiary biotic transition |journal=] |location=London |publisher=] |volume=154 |issue=2 |pages=265–292 |doi=10.1144/gsjgs.154.2.0265|bibcode=1997JGSoc.154..265M |s2cid=129654916 |issn=0016-7649}}</ref> Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs (such as '']'') could fly or at least glide, and others, such as ], had ] habits.<ref name=theropods/>
The hip joint arrangement described above allowed an erect stance, in which hind limbs were situated directly beneath the body or 'underslung'. This stance is like that of most mammals today but unlike that of other reptiles, which have a less erect posture and limbs splayed out to either side. The vertical action of the limbs in dinosaurs allowed for more efficient and faster locomotion, compared to the clumsier and slower movement of other 'sprawled' reptiles. It also allowed many types of dinosaurs to become bipedal.


===Distinguishing anatomical features===
]'' skeleton at the ] in ].]]
While recent discoveries have made it more difficult to present a universally agreed-upon list of their distinguishing features, nearly all dinosaurs discovered so far share certain modifications to the ancestral archosaurian skeleton, or are clearly descendants of older dinosaurs showing these modifications. Although some later groups of dinosaurs featured further modified versions of these traits, they are considered typical for Dinosauria; the earliest dinosaurs had them and passed them on to their descendants. Such modifications, originating in the most recent common ancestor of a certain taxonomic group, are called the ] of such a group.<ref name="B012"/>
'''Taxonomic definition'''
]'']]


A detailed assessment of archosaur interrelations by ]<ref name=nesbitt2011/> confirmed or found the following twelve unambiguous synapomorphies, some previously known:
Under ], dinosaurs are defined as all descendants of the most recent common ancestor of '']'' and modern ]. They are divided into ] (''bird-hipped'') and ] (''lizard-hipped''), depending upon ] structure. Ornithischian dinosaurs had a four-pronged pelvic configuration, incorporating a caudally-directed (rear-pointing) ] bone with (most commonly) a forward-pointing process. By contrast, the pelvic structure of saurischian dinosaurs was three-pronged, and featured a pubis bone directed cranially, or forwards, only. Ornithischia includes all ] sharing a more recent common ancestor with ''Triceratops'' than with Saurischia, while Saurischia includes those taxa sharing a more recent common ancestor with ''birds'' than with Ornithischia. It has also been suggested that Dinosauria be defined as all the descendants of the most recent common ancestor of '']'' and '']'', because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria.
* In the ], a supratemporal fossa (excavation) is present in front of the ], the main opening in the rear skull roof
* ], obliquely backward-pointing processes on the rear top corners of the anterior (front) neck ]e behind the ] and ], the first two neck vertebrae
* Apex of a deltopectoral crest (a projection on which the ] muscles attach) located at or more than 30% down the length of the ] (upper arm bone)
* ], a lower arm bone, shorter than 80% of humerus length
* ] (projection where the ] muscle attaches on the inner rear shaft) on the ] (thigh bone) is a sharp flange
* Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft
* On the ] and ], upper ankle bones, the proximal articular facet, the top connecting surface, for the ] occupies less than 30% of the transverse width of the element
* Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity, the inner space of the braincase
* In the pelvis, the proximal articular surfaces of the ] with the ] and the ] are separated by a large concave surface (on the upper side of the ischium a part of the open hip joint is located between the contacts with the pubic bone and the ilium)
* ] on the ] (protruding part of the top surface of the shinbone) arcs anterolaterally (curves to the front and the outer side)
* Distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia (the rear surface of the lower end of the shinbone)
* Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it touches the fibula, has a hollow profile)


Nesbitt found a number of further potential synapomorphies and discounted a number of synapomorphies previously suggested. Some of these are also present in ], which Nesbitt recovered as a sister group to Dinosauria, including a large anterior trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and pubis, the presence of a cnemial crest on the tibia and of an ascending process on the astragalus, and many others.<ref name=MJB04dino/>
There is an almost universal consensus among paleontologists that ]s are the descendants of ] dinosaurs. Using the strict ] definition that all descendants of a single common ancestor are related, modern birds ''are'' dinosaurs and dinosaurs are, therefore, not extinct. Modern ]s are classified by most paleontologists as belonging to the subgroup ], which are ]s, which are ], which are ], which are dinosaurs.


]s (sprawling), dinosaurs and ]s (erect), and ]ns (pillar-erect)]]
However, referring to birds as 'avian dinosaurs' and to all other dinosaurs as 'non-avian dinosaurs' is cumbersome. Birds are still referred to as birds, at least in popular usage and among ]s. It is also technically correct to refer to birds as a distinct group under the older ] system, which accepts ] taxa that exclude some descendants of a single common ancestor. Paleontologists mostly use ], which classifies birds as dinosaurs, but some biologists of the older generation do not.


A variety of other skeletal features are shared by dinosaurs. However, because they either are common to other groups of archosaurs or were not present in all early dinosaurs, these features are not considered to be synapomorphies. For example, as ]s, dinosaurs ancestrally had two pairs of ]e (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the ] and lower jaw.<ref name=TRHJ00/> Additionally, several characteristics once thought to be synapomorphies are now known to have appeared before dinosaurs, or were absent in the earliest dinosaurs and independently evolved by different dinosaur groups. These include an elongated ], or shoulder blade; a ] composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in '']'');<ref name=MJB04dino/> and a perforate ], or hip socket, with a hole at the center of its inside surface (closed in '']'', for example).<ref name="UC Berkeley Journal of Earth Sciences">{{cite web |url=https://ucmp.berkeley.edu/diapsids/dinomm.html |last1=Smith |first1=Dave |display-authors=et al. |title=Dinosauria: Morphology |publisher=] |location=Berkeley |access-date=October 16, 2019}}</ref><ref name=LARB99/> Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the ] epoch are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature.<ref name=NIP07/>
For clarity, this article will use 'dinosaur' as a synonym for 'non-avian dinosaur', and 'bird' as a synonym for 'avian dinosaur' (meaning any animal that evolved from the common ancestor of '']'' and modern birds). The term 'non-avian dinosaur' will be used for emphasis as needed. It should be noted that this article's definition of 'bird' differs from the definition common in everyday language; to most non-scientists, a 'bird' is simply a two-legged animal with wings and feathers.


Dinosaurs stand with their hind limbs erect in a manner similar to ], but distinct from most other reptiles, whose limbs sprawl out to either side.<ref name=Holland1909/> This posture is due to the development of a laterally facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur.<ref name=MJB00/> Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that ].<ref name=RC05/> Erect limbs probably also helped support the ] of large size by reducing bending stresses on limbs.<ref name=TKMB07/> Some non-dinosaurian archosaurs, including ]ns, also had erect limbs but achieved this by a "pillar-erect" configuration of the hip joint, where instead of having a projection from the femur insert on a socket on the hip, the ] was rotated to form an overhanging shelf.<ref name=TKMB07/>
===Size===
While the evidence is incomplete, it is clear that, as a group, dinosaurs were large. Even by dinosaur standards, the ]s were gigantic. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest were an ] more massive than anything else that has since walked the Earth. Giant prehistoric ]s such as the '']'' and the Columbian ] were dwarfed by the giant sauropods, and only a handful of modern aquatic animals approach them in size &mdash; most notably the ], which reaches up to 190,000 kg (209 tons) and 33.5 m (110 ft) in length.


==History of study==
Most dinosaurs, however, were much smaller than the giant sauropods. Current evidence suggests that dinosaur average size varied through the Triassic, early Jurassic, late Jurassic and Cretaceous periods.<ref> Working hypothesis for body size.</ref> According to paleontologist Bill Erickson, estimates of median dinosaur weight range from 500 kg to 5 ]s; a recent study of 63 dinosaur genera yielded an average weight in excess of 850 kg &mdash; comparable to the weight of a grizzly bear &mdash; and a median weight of nearly 2 tons, or about as much as a giraffe. This contrasts sharply with the size of modern mammals; on average, mammals weigh only 863 grams, or about as much as a large rodent. The smallest dinosaur was bigger than two-thirds of all current mammals; the majority of dinosaurs were bigger than all but 2% of living mammals.<ref> Soruce of Erickson quote.</ref>
{{Further|History of paleontology}}


===Pre-scientific history===
]'', outside the ].]]
Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese considered them to be ] bones and documented them as such. For example, '']''&nbsp;({{zh|t=華陽國志|labels=no}}), a ] compiled by ]&nbsp;({{zh|t=常璩|labels=no}}) during the ] (265–316), reported the discovery of dragon bones at Wucheng in ] Province.<ref name=dong1992/> Villagers in central ] have long unearthed fossilized "dragon bones" for use in ].<ref name=BBCdinobonemed/> In ], dinosaur fossils were generally believed to be the remains of ]s and other ] creatures.<ref name=benton2000>{{harvnb|Paul|2000|pp=10–44|loc=chpt. 1: "A Brief History of Dinosaur Paleontology" by Michael J. Benton.}}</ref>


===Early dinosaur research===
'''Largest and smallest dinosaurs'''
]]]


Scholarly descriptions of what would now be recognized as dinosaur bones first appeared in the late 17th century in England. Part of a bone, now known to have been the femur of a '']'',<ref name=WAS97/> was recovered from a limestone quarry at ] near ], Oxfordshire, in 1676. The fragment was sent to ], Professor of Chemistry at the ] and first curator of the ], who published a description in his ''The Natural History of Oxford-shire'' (1677).<ref>{{harvnb|Plot|1677|pp=131–139; illus. opp. p. 142, fig. 4}}</ref> He correctly identified the bone as the lower extremity of the femur of a large animal, and recognized that it was too large to belong to any known species. He therefore concluded it to be the femur of a huge human, perhaps a ] or another type of giant featured in legends.<ref>{{harvnb|Plot|1677|p=}}</ref><ref>{{cite web |url=http://www.oum.ox.ac.uk/learning/pdfs/plot.pdf |title=Robert Plot |author=<!--Staff writer(s); no by-line.--> |year=2006 |website=Learning more |publisher=] |location=Oxford |archive-url=https://web.archive.org/web/20061001094736/http://www.oum.ox.ac.uk/learning/pdfs/plot.pdf |archive-date=October 1, 2006 |access-date=November 14, 2019}}</ref> ], a friend of ], published ''Lithophylacii Britannici ichnographia'' (1699), the first scientific treatment of what would now be recognized as a dinosaur. In it he described and named a sauropod ], "]",<ref name=L99/><ref name=DS02/> that had been found in Caswell, near ], Oxfordshire.<ref name=G45/>
Only a tiny percentage of animals ever fossilize, and most of these remain buried in the earth. Few of the specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork. As a result, scientists will probably never be certain of the ].


]'s coining of the word ''dinosaur'', in the 1842 revised version of his talk at an 1841 meeting of the ].]]
]''.]]
The tallest and heaviest dinosaur known from a complete skeleton is the '']'' specimen that was discovered in ] between 1907&ndash;12. It is now mounted and on display at the ] of ] and is 12 m (38 ft) tall and probably weighed between 30,000–60,000 kg (33&ndash;66 ]s). The longest complete dinosaur is the 27 m (89 ft) long '']'', which was discovered in ] in the ] and displayed in ]'s ] in 1907.


Between 1815 and 1824, the Rev ], the first Reader of Geology at the University of Oxford, collected more fossilized bones of ''Megalosaurus'' and became the first person to describe a non-avian dinosaur in a ].<ref name=WAS97/><ref name=buckland1824/> The second non-avian dinosaur genus to be identified, '']'', was purportedly discovered in 1822 by ], the wife of English geologist ], though this is disputed and some historians say Gideon had acquired remains years earlier. Gideon Mantell recognized similarities between his fossils and the bones of modern ]s and published his findings in 1825.<ref name=GM25/><ref name=HDS97/>
There were larger dinosaurs, but knowledge of them is based entirely on a small number of incomplete fossil samples. The largest ] specimens on record were all discovered in the 1970s or later, and include the massive '']'', which may have weighed 80,000–100,000 kg (88&ndash;121 tons); the longest, the 40 m (130 ft) long '']''; and the tallest, the 18 m (60 ft) '']'', which could have reached a sixth-floor window. The largest known ] dinosaur was '']'', reaching a length of 16-18 meters (53-60 ft), and weighing in at 9 tons. Other large meat-eaters included '']'', '']'', '']'' and '']''.
Not including modern birds like the ], the smallest dinosaurs known were about the size of a ] or a ]. The theropods '']'', '']'', and '']'' were all under 60 cm (2 ft) in length.


The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English paleontologist Sir Richard Owen coined the term "dinosaur", using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world.<ref name=":2" /><ref name=":3" /><ref name=":4" /><ref name=Owen1841/><ref>{{cite Merriam-Webster|Dinosauria|access-date=November 10, 2019}}</ref> The term is derived {{ety|grc|'']'' (deinos)|terrible, potent or fearfully great||'']'' (sauros)|lizard or reptile}}.<ref name=Owen1841/><ref name=LSJ/> Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it also to evoke their size and majesty.<ref name=FBS97/> Owen recognized that the remains that had been found so far, ''Iguanodon'', ''Megalosaurus'' and '']'', shared distinctive features, and so decided to present them as a distinct taxonomic group. As clarified by British geologist and historian Hugh Torrens, Owen had given a presentation about fossil reptiles to the British Association for the Advancement of Science in 1841, but reports of the time show that Owen did not mention the word "dinosaur", nor recognize dinosaurs as a distinct group of reptiles in his address. He introduced the Dinosauria only in the revised text version of his talk published in April 1842.<ref name=":2" /><ref name=":3" /> With the backing of ], the husband of ], Owen established the ], to display the national collection of dinosaur fossils and other biological and geological exhibits.<ref name=owen94>{{harvnb|Rupke|1994}}</ref>
===Behavior===
]'' was discovered in 1978.]]
Interpretations of dinosaur behavior are generally based on the pose of body fossils and their ], ]s of their ], and comparisons with modern animals in similar ]s. As such, the current understanding of dinosaur behavior relies on speculation, and will likely remain controversial for the foreseeable future. However, there is general agreement that some behaviors which are common in crocodiles and birds, dinosaurs' closest living relatives, were also common among dinosaurs.


===Discoveries in North America===
The first perceived direct evidence of ]ing behavior was the 1878 discovery of 31 '']'' dinosaurs which were thought to have perished together in ], ], after they fell into a deep, flooded ] and drowned.<ref>Johan Yans, Jean Dejax, Denise Pons, Christian Dupuis, Philippe Taquet.(2005) Palaeontological and geodynamical implications of the palynological dating of the wealden facies sediments of Bernissart (Mons Basin, Belgium). ''C. R. Palevol 4 (2005)'' 135–150.</ref> Despite the deposition of those skeletons being now regarded as more gradual,<ref>Deposition of ''Iguanodon'' skeletons ocurred in at least 3 different events.</ref> other, well supported, mass death sites were subsequently discovered. Those, along with multiple trackways, suggest that ] or ] behavior was common in many dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that ]s (hadrosaurids) may have moved in great herds, like the ] or the African ]. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in ], England,<ref>Day, J.J. and Upchurch, P. (2002). Sauropod Trackways, Evolution, and Behavior. ''Science'' 296:1659. </ref> and others kept their young in the middle of the herd for defense according to trackways at Davenport Ranch, ]. Dinosaurs may have congregated in herds for defense, for ] purposes, or to provide protection for their young.
{{multiple image
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In 1858, ] discovered the first known American dinosaur, in ] pits in the small town of ]. (Although fossils had been found before, their nature had not been correctly discerned.) The creature was named '']''. It was an extremely important find: ''Hadrosaurus'' was one of the first nearly complete dinosaur skeletons found (] was in 1834, in ]), and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of interests in dinosaurs in the United States, known as dinosaur mania.<ref name=weishampel06>{{cite journal |last1=Prieto-Marquez |first1=Albert |last2=Weishampel |first2=David B. |last3=Horner |first3=John R. |date=March 2006 |title=The dinosaur ''Hadrosaurus foulkii'', from the Campanian of the East Coast of North America, with a reevaluation of the genus |url=https://app.pan.pl/archive/published/app51/app51-077.pdf |url-status=live |journal=Acta Palaeontologica Polonica |location=Warsaw |publisher=Institute of Paleobiology, Polish Academy of Sciences |volume=51 |issue=1 |pages=77–98 |issn=0567-7920 |archive-url=https://web.archive.org/web/20190622113255/https://app.pan.pl/archive/published/app51/app51-077.pdf |archive-date=June 22, 2019 |access-date=November 5, 2019}}</ref>
]'s 1978 discovery of a '']'' ("good mother dinosaur") ]ing ground in ] demonstrated that parental care continued long after birth among the ]s.<ref>Lessem, D. and Glut, D.F. (1993). ''The Dinosaur Society's Dinosaur Encyclopedia''. Random House Inc. ISBN 0-679-41770-2. </ref><ref> A juvenile ] skeleton was found.</ref> There is also evidence that other Cretaceous-era dinosaurs, like the ]n sauropod '']'' (1997 discovery), had similar nesting behaviors, and that the animals congregated in huge nesting colonies like those of ]s. The ]n ]n '']'' was discovered in a ]-like ]ing position in 1993, which may mean it was covered with an insulating layer of feathers that kept the ] warm.<ref> ] nests or ]?</ref> Trackways have also confirmed parental behavior among sauropods and ornithopods from the ] in northwestern ].<ref> Footprints show maternal instinct after leaving the nest.</ref> Nests and eggs have been found for most major groups of dinosaurs, and it appears likely that dinosaurs communicated with their young, in a manner similar to modern birds and crocodiles.


Dinosaur mania was exemplified by the fierce rivalry between ] and ], both of whom raced to be the first to find new dinosaurs in what came to be known as the ]. This fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their diggers often used ] to unearth bones. Modern paleontologists would find such methods crude and unacceptable, since blasting easily destroys fossil and stratigraphic evidence. Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast: Marsh unearthed 86 new species of dinosaur and Cope discovered 56, a total of 142 new species. Cope's collection is now at the ] in New York City, while Marsh's is at the ] at ].<ref name=Holmes/>
The ]s and frills of some dinosaurs, like the ]ns, ]s and ]es, may have been too fragile to be used for active defense, so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and ]. The nature of dinosaur ] also remains enigmatic, and is an active area of research. For example, recent evidence suggests that the hollow crests of the lambeosaurines may have functioned as ]s used for a wide range of ]s.


==="Dinosaur renaissance" and beyond===
From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the ] in 1971. It included a '']'' attacking a '']'',<ref> The discovery of two ] dinosaurs entangled together proved many theories.</ref> proving that dinosaurs did indeed attack and eat each other. While ]istic behavior among ]s is no surprise,<ref> The mystery of a dinosaur ].</ref> this too was confirmed by tooth marks from Madagascar in 2003.<ref>Rogers, R.R., Krause, D.W. and Rogers, K.C. (2003). Cannibalism in the Madagascan dinosaur Majungatholus atopus. ''Nature'' 422:515-518..</ref>
{{Main|Dinosaur renaissance}}
]'s original restoration of '']'', published in 1969]]
] caused a pause in palaeontological research; after the war, research attention was also diverted increasingly to fossil mammals rather than dinosaurs, which were seen as sluggish and cold-blooded.<ref name="taylor2010">{{cite journal |last1=Taylor |first1=M.P. |title=Sauropod dinosaur research: a historical review |journal=Geological Society, London, Special Publications |year=2010 |volume=343 |issue=1 |pages=361–386 |doi=10.1144/SP343.22|bibcode=2010GSLSP.343..361T |s2cid=910635 }}</ref><ref name="Naish">{{cite book |last=Naish |first=D. |year=2009 |title=The Great Dinosaur Discoveries |pages=89–93 |publisher=A & C Black Publishers Ltd. |location=London, UK |isbn=978-1-4081-1906-8}}</ref> At the end of the 1960s, however, the field of dinosaur research experienced a surge in activity that remains ongoing.<ref name="arbour2018">{{cite journal |last1=Arbour |first1=V. |year=2018 |title=Results roll in from the dinosaur renaissance |journal=Science |volume=360 |issue=6389 |pages=611 |doi=10.1126/science.aat0451|bibcode=2018Sci...360..611A |s2cid=46887409 }}</ref> Several seminal studies led to this activity. First, ] discovered the bird-like ] theropod '']'' and described it in 1969. Its anatomy indicated that it was an active predator that was likely warm-blooded, in marked contrast to the then-prevailing image of dinosaurs.<ref name="taylor2010"/> Concurrently, ] published a series of studies that likewise argued for active lifestyles in dinosaurs based on anatomical and ecological evidence (see {{section link||Physiology}}),<ref name="bakker1968"/><ref name="bakker1972"/> which were subsequently summarized in his 1986 book '']''.<ref name=bakker86>{{harvnb|Bakker|1986}}</ref>


] with a mounted skeleton of a ] ('']'')]]
There seem to have been no burrowing species of dinosaur and few climbing species. This is somewhat surprising when compared to the later mammalian radiation in the ], which included many species of these types. As to how the animals moved, ] has provided significant insight. For example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have demonstrated how fast dinosaurs could run,<ref> Gait and his formula on estimating a dinosaur's speed.</ref><ref> More on Gait and his speed calculations.</ref> whether ]s could create ]s via ]-like tail snapping,<ref>Douglas, K. and Young, S. (1998). The dinosaur detectives. ''New Scientist'' 2130:24. .</ref> whether giant theropods had to slow down when rushing for food to avoid fatal injuries,<ref>Hecht, J. (1998). The deadly dinos that took a dive. ''New Scientist'' 2130. .</ref> and if sauropods could float.<ref>Henderson, D.M. (2003). Effects of stomach stones on the buoyancy and equilibrium of a floating crocodilian: A computational analysis. ''Canadian Journal of Zoology'' 81:1346-1357. .</ref>
New revelations were supported by an increase in dinosaur discoveries. Major new dinosaur discoveries have been made by paleontologists working in previously unexplored regions, including India, South America, Madagascar, Antarctica, and most significantly China. Across theropods, sauropodomorphs, and ornithischians, the number of named genera began to increase exponentially in the 1990s.<ref name="Genera900"/> In 2008 over 30 new species of dinosaurs were named each year.<ref name="benton2008"/> At least sauropodomorphs experienced a further increase in the number of named species in the 2010s, with an average of 9.3 new species having been named each year between 2009 and 2020. As a consequence, more sauropodomorphs were named between 1990 and 2020 than in all previous years combined.<ref name="cashmore2020">{{cite journal |last1=Cashmore |first1=D.D. |last2=Mannion |first2=P.D. |last3=Upchurch |first3=P. |last4=Butler |first4=R.J. |year=2020 |title=Ten more years of discovery: revisiting the quality of the sauropodomorph dinosaur fossil record |journal=Palaeontology |volume=63 |issue=6 |pages=951–978 |doi=10.1111/pala.12496|bibcode=2020Palgy..63..951C |s2cid=219090716 |doi-access=free }}</ref> These new localities also led to improvements in overall specimen quality, with new species being increasingly named not on scrappy fossils but on more complete skeletons, sometimes from multiple individuals. Better specimens also led to new species being invalidated less frequently.<ref name="benton2008">{{cite journal |last1=Benton |first1=M.J. |year=2008 |title=Fossil quality and naming dinosaurs |journal=Biology Letters |volume=4 |issue=6 |pages=729–732 |doi=10.1098/rsbl.2008.0402|pmid=18796391 |pmc=2614166 }}</ref> Asian localities have produced the most complete theropod specimens,<ref name="cashmore2019">{{cite journal |last1=Cashmore |first1=D.D. |last2=Butler |first2=R.J. |year=2019 |title=Skeletal completeness of the non-avian theropod dinosaur fossil record |journal=Palaeontology |volume=62 |issue=6 |pages=951–981 |doi=10.1111/pala.12436|s2cid=197571209 |doi-access=free |bibcode=2019Palgy..62..951C }}</ref> while North American localities have produced the most complete sauropodomorph specimens.<ref name="cashmore2020"/>


Prior to the dinosaur renaissance, dinosaurs were mostly classified using the traditional rank-based system of ]. The renaissance was also accompanied by the increasingly widespread application of ], a more objective method of classification based on ancestry and shared traits, which has proved tremendously useful in the study of dinosaur systematics and evolution. Cladistic analysis, among other techniques, helps to compensate for an often incomplete and fragmentary fossil record.<ref name="holtz1997">{{cite book |last1=Holtz |first1=T.R. Jr. |last2=Brett-Surman |first2=M.K. |year=1997 |chapter=The Taxonomy and Systematics of Dinosaurs |title=The Complete Dinosaur |publisher=Indiana University Press |location=Bloomington |pages=209–223 |isbn=978-0-253-33349-0 |chapter-url=https://books.google.com/books?id=Hk5ecvEv0GcC&pg=PA209}}</ref><ref name="NYT-20161208">{{cite news |last=St. Fleur |first=Nicholas |title=That Thing With Feathers Trapped in Amber? It Was a Dinosaur Tail |url=https://www.nytimes.com/2016/12/08/science/dinosaur-feathers-amber.html |url-status=live |url-access=registration |date=December 8, 2016 |department=Trilobites |work=The New York Times |location=New York |issn=0362-4331 |archive-url=https://web.archive.org/web/20170831181949/https://www.nytimes.com/2016/12/08/science/dinosaur-feathers-amber.html |archive-date=August 31, 2017 |access-date=December 8, 2016}}</ref> Reference books summarizing the state of dinosaur research, such as ] and colleagues' '']'', made knowledge more accessible<ref name="lockley2000">{{cite journal |last1=Lockley |first1=M.G. |last2=Wright |first2=J.L. |year=2000 |title=Reading About Dinosaurs – An Annotated Bibliography of Books |journal=Journal of Geoscience Education |volume=48 |issue=2 |pages=167–178 |doi=10.5408/1089-9995-48.2.167|bibcode=2000JGeEd..48..167L |s2cid=151426669 }}</ref> and spurred further interest in dinosaur research. The release of the first and second editions of ''The Dinosauria'' in 1990 and 2004, and of a review paper by ] in 1998, were accompanied by increases in the number of published ]s for dinosaurs.<ref name="lloyd2008">{{cite journal |last1=Lloyd |first1=G.T. |last2=Davis |first2=K.E. |last3=Pisani |first3=D. |last4=Tarver |first4=J.E. |last5=Ruta |first5=R. |last6=Sakamoto |first6=M. |last7=Hone |first7=D.W.E. |last8=Jennings |first8=R. |last9=Benton |first9=M.J. |year=2008 |title=Dinosaurs and the Cretaceous Terrestrial Revolution |journal=Proceedings of the Royal Society B |volume=275 |issue=1650 |pages=2483–2490 |doi=10.1098/rspb.2008.0715|pmid=18647715 |pmc=2603200 }}</ref>
==Evolution of dinosaurs==
]'', an early dinosaur.]]
Dinosaurs diverged from their ] ancestors approximately 230 million years ago during the Middle to Late ] period, roughly 20 million years after the ] wiped out an estimated 95% of all life on Earth.<ref>Citation for Permian/Triassic extinction event, percentage of animal species that went extinct. </ref><ref>Another citation for P/T event data. </ref> ] of fossils from the early dinosaur ] '']'' establishes its presence in the fossil record at this time. Paleontologists believe ''Eoraptor'' resembles the ] of all dinosaurs;<ref>Hayward, T. (1997). The First Dinosaurs. ''Dinosaur Cards''. Orbis Publishing Ltd. D36040612.</ref> if this is true, its traits suggest that the first dinosaurs were small, bipedal predators.<ref>Sereno, P.C., C.A. Forster, R.R. Rogers, and A.M. Monetta. 1993. Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria. Nature 361:64-66.</ref> Such a notion is corroborated by the more primitive dinosaur-like ornithodirans hailing from ] strata of Argentina such as '']'' and '']'', which were also small bipedal, perhaps leaping, predators.


===Soft tissue and molecular preservation===
The first few lines of primitive dinosaurs ] through the rest of the Triassic period; dinosaur species quickly evolved the specialized features and range of sizes needed to exploit nearly every terrestrial ]. During the period of dinosaur predominance, which encompassed the ensuing ] and ] periods, nearly every known land animal larger than 1 meter in length was a dinosaur.
]'' specimen's skin impressions found in 1999]]
Dinosaur fossils are not limited to bones, but also include imprints or mineralized remains of skin coverings, organs, and other tissues. Of these, skin coverings based on ] proteins are most easily preserved because of their ]ed, ] molecular structure.<ref name="schweitzer2011">{{cite journal |last1=Schweitzer |first1=M.H. |year=2011 |title=Soft Tissue Preservation in Terrestrial Mesozoic Vertebrates |journal=Annual Review of Earth and Planetary Sciences |volume=39 |pages=187–216 |doi=10.1146/annurev-earth-040610-133502|bibcode=2011AREPS..39..187S }}</ref> Fossils of keratin-based skin coverings or bony skin coverings are known from most major groups of dinosaurs. Dinosaur fossils with scaly skin impressions have been found since the 19th century. ] discovered a sauropod forelimb with preserved skin in 1852 that was incorrectly attributed to a crocodile; it was correctly attributed by Marsh in 1888 and subject to further study by ] in 1917.<ref name="hooley1917"/> Among ornithischians, in 1884 Jacob Wortman found skin impressions on the first known specimen of '']'', which were largely destroyed during the specimen's excavation.<ref name="osborn1912">{{cite journal |first1=H.F. |last1=Osborn |year=1912 |title=Integument of the iguanodont dinosaur ''Trachodon'' |journal=Memoirs of the American Museum of Natural History |volume=1 |pages=33–54}}</ref> Owen and Hooley subsequently described skin impressions of '']'' and ''Iguanodon'' in 1885 and 1917.<ref name="hooley1917">{{cite journal |last1=Hooley |first1=R.W. |year=1917 |title=II—On the Integument of ''Iguanodon bernissartensis'', Boulenger, and of ''Morosaurus becklesii'', Mantell |journal=Geological Magazine |volume=4 |issue=4 |pages=148–150 |doi=10.1017/s0016756800192386|bibcode=1917GeoM....4..148H |s2cid=129640665 |url=https://zenodo.org/record/1537494 }}</ref> Since then, scale impressions have been most frequently found among hadrosaurids, where the impressions are known from nearly the entire body across multiple specimens.<ref name="bell2014">{{cite book |last1=Bell |first1=P.R. |year=2014 |chapter=A review of hadrosaur skin impressions |editor-last1=Eberth |editor-first1=D. |editor-last2=Evans |editor-first2=D. |title=The Hadrosaurs: Proceedings of the International Hadrosaur Symposium |pages=572–590 |location=Bloomington |publisher=Princeton University Press}}</ref>


{{multiple image|align=left|perrow=1/2|caption_align=center
The ], which occurred approximately 65 million years ago at the end of the Cretaceous period, caused the extinction of all dinosaurs except for the line that had already given rise to the first birds. Other ] species related to the dinosaurs also survived the event.
|image1=Sinosauropteryx color.jpg|caption1=Color restoration of '']''
|image2=Psittacosaurus (Vinther et al. 2016, cropped).png|caption2=Color restoration of '']''
}}
Starting from the 1990s, major discoveries of exceptionally preserved fossils in deposits known as conservation ]n contributed to research on dinosaur soft tissues.<ref name="eliason2017">{{cite journal |last1=Eliason |first1=C.M. |last2=Hudson |first2=L. |last3=Watts |first3=T. |last4=Garza |first4=H. |last5=Clarke |first5=J.A. |year=2017 |title=Exceptional preservation and the fossil record of tetrapod integument |journal=Proceedings of the Royal Society B |volume=284 |issue=1862 |pages=1–10 |doi=10.1098/rspb.2017.0556|pmid=28878057 |pmc=5597822 }}</ref><ref name="benton1998">{{cite journal |last1=Benton |first1=M.J. |year=1998 |title=Dinosaur fossils with soft parts |journal=Trends in Ecology & Evolution |volume=13 |issue=8 |pages=303–304 |doi=10.1016/s0169-5347(98)01420-7|pmid=21238317 |bibcode=1998TEcoE..13..303B |url=http://doc.rero.ch/record/14715/files/PAL_E1646.pdf }}</ref> Chiefly among these were the rocks that produced the ] (Early Cretaceous) and ] (Mid-to-Late Jurassic) ]s of northeastern China, from which hundreds of dinosaur specimens bearing impressions of feather-like structures (both closely related to birds and otherwise, see {{section link||Origin of birds}}) have been described by ] and colleagues.<ref name="zhou2017">{{cite journal |last1=Zhou |first1=Z.-H. |last2=Wang |first2=Y. |year=2017 |title=Vertebrate assemblages of the Jurassic Yanliao Biota and the Early Cretaceous Jehol Biota: Comparisons and implications |journal=Palaeoworld |volume=26 |issue=2 |pages=241–252 |doi=10.1016/j.palwor.2017.01.002}}</ref><ref name="norell2005">{{cite journal |last1=Norell |first1=M.A. |last2=Xu |first2=X. |year=2005 |title=Feathered Dinosaurs |journal=Annual Review of Earth and Planetary Sciences |volume=33 |pages=277–299 |doi=10.1146/annurev.earth.33.092203.122511|bibcode=2005AREPS..33..277N }}</ref> In living reptiles and mammals, pigment-storing cellular structures known as ]s are partially responsible for producing colouration.<ref name="roy2020"/><ref name="vinther2020">{{cite journal |last1=Vinther |first1=J. |year=2020 |title=Reconstructing Vertebrate Paleocolor |journal=Annual Review of Earth and Planetary Sciences |volume=48 |pages=345–375 |doi=10.1146/annurev-earth-073019-045641|bibcode=2020AREPS..48..345V |s2cid=219768255 |doi-access=free }}</ref> Both chemical traces of ] and characteristically shaped melanosomes have been reported from feathers and scales of Jehol and Yanliao dinosaurs, including both theropods and ornithischians.<ref name="zhang2010">{{cite journal |last1=Zhang |first1=F. |last2=Kearns |first2=S.L. |last3=Orr |first3=P.J. |last4=Benton |first4=M.J. |last5=Zhou |first5=Z. |last6=Johnson |first6=D. |last7=Xu |first7=X. |last8=Wang |first8=X. |year=2010 |title=Fossilized melanosomes and the colour of Cretaceous dinosaurs and birds |journal=Nature |volume=463 |issue=7284 |pages=1075–1078 |doi=10.1038/nature08740 |pmid=20107440 |bibcode=2010Natur.463.1075Z |s2cid=205219587 |url=http://oro.open.ac.uk/22432/2/41064696.pdf }}</ref> This has enabled multiple full-body reconstructions of ], such as for '']''<ref name="smithwick2017">{{cite journal |last1=Smithwick |first1=F.M. |last2=Nicholls |first2=R. |last3=Cuthill |first3=I.C. |last4=Vinther |first4=J. |year=2017 |title=Countershading and Stripes in the Theropod Dinosaur ''Sinosauropteryx'' Reveal Heterogeneous Habitats in the Early Cretaceous Jehol Biota |journal=Current Biology |volume=27 |issue=21 |pages=3337–3343.e2 |doi=10.1016/j.cub.2017.09.032 |pmid=29107548 |doi-access=free|bibcode=2017CBio...27E3337S |hdl=1983/8ee95b15-5793-42ad-8e57-da6524635349 |hdl-access=free }}</ref> and '']''<ref name="vinther2016">{{cite journal |last1=Vinther |first1=J. |last2=Nicholls |first2=R. |last3=Lautenschlager |first3=S. |last4=Pittman |first4=M. |last5=Kaye |first5=T.G. |last6=Rayfield |first6=E. |last7=Mayr |first7=G. |last8=Cuthill |first8=I.C. |year=2016 |title=3D Camouflage in an Ornithischian Dinosaur |journal=Current Biology |volume=26 |issue=18 |pages=2456–2462 |doi=10.1016/j.cub.2016.06.065 |pmid=27641767 |pmc=5049543|bibcode=2016CBio...26.2456V }}</ref> by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur fossils from other localities.<ref name="roy2020">{{cite journal |last1=Roy |first1=A. |last2=Pittman |first2=M. |last3=Saitta |first3=E.T. |last4=Kaye |first4=T.G. |last5=Xu |first5=X. |year=2020 |title=Recent advances in amniote palaeocolour reconstruction and a framework for future research |journal=Biological Reviews |volume=95 |issue=1 |pages=22–50 |doi=10.1111/brv.12552|pmid=31538399 |pmc=7004074 }}</ref> (However, some researchers have also suggested that fossilized melanosomes represent bacterial remains.<ref name="lindgren2015">{{cite journal |last1=Lindgren |first1=J. |last2=Moyer |first2=A. |last3=Schweitzer |first3=M.H. |last4=Sjövall |first4=P. |last5=Uvdal |first5=P. |last6=Nilsson |first6=D.E. |last7=Heimdal |first7=J. |last8=Engdahl |first8=A. |last9=Gren |first9=J.A. |last10=Schultz |first10=B.P. |last11=Kear |first11=B.P. |year=2015 |title=Interpreting melanin-based coloration through deep time: a critical review |journal=Proceedings of the Royal Society B |volume=282 |issue=1813 |pages=20150614 |pmid=26290071 |doi=10.1098/rspb.2015.0614 |pmc=4632609}}</ref><ref name="schweitzer2015">{{cite journal |last1=Schweitzer |first1=M.H. |last2=Lindgren |first2=J. |last3=Moyer |first3=A.E. |year=2015 |title=Melanosomes and ancient coloration re-examined: a response to Vinther 2015 (DOI 10.1002/bies.201500018) |journal=BioEssays |volume=37 |issue=11 |pages=1174–1183 |doi=10.1002/bies.201500061|pmid=26434749 |s2cid=45178498 }}</ref>) Stomach contents in some Jehol and Yanliao dinosaurs closely related to birds have also provided indirect indications of diet and digestive system anatomy (e.g., ]).<ref name="zhou2014">{{cite journal |last1=Zhou |first1=Z. |year=2014 |title=The Jehol Biota, an Early Cretaceous terrestrial Lagerstätte: new discoveries and implications |journal=National Science Review |volume=1 |issue=4 |pages=543–559 |doi=10.1093/nsr/nwu055|doi-access=free }}</ref><ref name="oconnor2019">{{cite journal |last1=O'Connor |first1=J.K. |last2=Zhou |first2=Z. |year=2019 |title=The evolution of the modern avian digestive system: insights from paravian fossils from the Yanliao and Jehol biotas |journal=Palaeontology |volume=63 |issue=1 |pages=13–27 |doi=10.1111/pala.12453|s2cid=210265348 }}</ref> More concrete evidence of internal anatomy has been reported in '']'' from the ] of Italy. It preserves portions of the intestines, colon, liver, muscles, and windpipe.<ref name="softtissue"/>


]'' fossil with intestines, ]]]
==Study of dinosaurs==
Concurrently, a line of work led by ], ], and colleagues reported various occurrences of preserved soft tissues and proteins within dinosaur bone fossils. Various mineralized structures that likely represented ]s and ] fibres had been found by Schweitzer and others in ] bones as early as 1991.<ref name="morell1993">{{cite journal |last1=Morell |first1=V. |year=1993 |title=Dino DNA: the Hunt and the Hype |journal=Science |volume=261 |issue=5118 |pages=160–162 |doi=10.1126/science.8327889|pmid=8327889 |bibcode=1993Sci...261..160M }}</ref><ref name="pawlicki1996">{{cite journal |last1=Pawlicki |first1=R. |last2=Korbel |first2=A. |last3=Kubiak |first3=H. |year=1996 |title=Cells, Collagen Fibrils and Vessels in Dinosaur Bone |journal=Nature |volume=211 |issue=5049 |pages=655–657 |doi=10.1038/211655a0|pmid=5968744 |s2cid=4181847 }}</ref><ref name="pawlicki1998">{{cite journal |last1=Pawlicki |first1=R. |last2=Nowogrodzka-Zagórska |first2=M. |year=1998 |title=Blood vessels and red blood cells preserved in dinosaur bones |journal=Annals of Anatomy - Anatomischer Anzeiger |volume=180 |issue=1 |pages=73–77 |doi=10.1016/S0940-9602(98)80140-4|pmid=9488909 }}</ref> However, in 2005, Schweitzer and colleagues reported that a femur of '']'' preserved soft, flexible tissue within, including ]s, ], and connective tissue (bone fibers) that had retained their microscopic structure.<ref name=Schweitzer2005/> This discovery suggested that original soft tissues could be preserved over geological time,<ref name="schweitzer2011"/> with multiple mechanisms having been proposed.<ref name="anderson2023">{{cite journal |last1=Anderson |first1=L.A. |year=2023 |title=A chemical framework for the preservation of fossil vertebrate cells and soft tissues |journal=Earth-Science Reviews |volume=240 |pages=104367 |doi=10.1016/j.earscirev.2023.104367|bibcode=2023ESRv..24004367A |s2cid=257326012 |doi-access=free }}</ref> Later, in 2009, Schweitzer and colleagues reported that a '']'' femur preserved similar microstructures, and ] techniques (based on ] binding) demonstrated the presence of proteins such as collagen, ], and ].<ref name="schweitzer2009">{{cite journal |last1=Schweitzer |first1=M.H. |last2=Zheng |first2=W. |last3=Organ |first3=C.L. |last4=Avci |first4=R. |last5=Suo |first5=Z. |last6=Freimark |first6=L.M. |last7=LeBleu |first7=V.S. |last8=Duncan |first8=M.B. |last9=van der Heiden |first9=M.G. |last10=Neveu |first10=J.M. |last11=Lane |first11=W.S. |last12=Cottrell |first12=J.S. |last13=Horner |first13=J.R. |last14=Cantley |first14=L.C. |last15=Kalluri |first15=R. |last16=Asara |first16=J.M. |year=2009 |title=Biomolecular characterization and protein sequences of the Campanian hadrosaur ''B. canadensis'' |journal=Science |volume=324 |issue=5927 |pages=626–631 |doi=10.1126/science.1165069|pmid=19407199 |bibcode=2009Sci...324..626S |s2cid=5358680 }}</ref> Both specimens yielded collagen protein sequences that were viable for ], which grouped them with birds as would be expected.<ref name="schweitzer2009"/><ref name="organ2008">{{cite journal |last1=Organ |first1=C.L. |last2=Schweitzer |first2=M.H. |last3=Zheng |first3=W. |last4=Freimark |first4=L.M. |last5=Cantley |first5=L.C. |last6=Asara |first6=J.M. |year=2008 |title=Molecular Phylogenetics of Mastodon and ''Tyrannosaurus rex'' |journal=Science |volume=320 |issue=5875 |pages=499 |doi=10.1126/science.1154284|pmid=18436782 |bibcode=2008Sci...320..499O |s2cid=24971064 }}</ref> The extraction of fragmentary DNA has also been reported for both of these fossils,<ref name="schweitzer2013">{{cite journal |last1=Schweitzer |first1=M.H. |last2=Zheng |first2=W. |last3=Cleland |first3=T.P. |last4=Bern |first4=M. |year=2013 |title=Molecular analyses of dinosaur osteocytes support the presence of endogenous molecules |journal=] |location=Amsterdam |publisher=Elsevier |volume=52 |issue=1 |pages=414–423 |doi=10.1016/j.bone.2012.10.010 |issn=8756-3282 |pmid=23085295}}</ref> along with a specimen of '']''.<ref name="bailleul2020">{{cite journal |last1=Bailleul |first1=A.M. |last2=Zheng |first2=W. |last3=Horner |first3=J.R. |last4=Hall |first4=B.K. |last5=Holliday |first5=C.M. |last6=Schweitzer |first6=M.H. |year=2020 |title=Evidence of proteins, chromosomes and chemical markers of DNA in exceptionally preserved dinosaur cartilage |journal=National Science Review |volume=7 |issue=4 |pages=815–822 |doi=10.1093/nsr/nwz206|pmid=34692099 |pmc=8289162 }}</ref> In 2015, Sergio Bertazzo and colleagues reported the preservation of collagen fibres and red blood cells in eight Cretaceous dinosaur specimens that did not show any signs of exceptional preservation, indicating that soft tissue may be preserved more commonly than previously thought.<ref name="bertazzo2015">{{cite journal |last1=Bertazzo |first1=S. |last2=Maidment |first2=S.C.R. |author2-link=Susannah Maidment |last3=Kallepitis |first3=C. |last4=Fearn |first4=S. |last5=Stevens |first5=M.M. |last6=Xie |first6=H.-N. |display-authors=3 |year=2015 |title=Fibres and cellular structures preserved in 75-million-year-old dinosaur specimens |journal=] |volume=6 |page=7352 |bibcode=2015NatCo...6.7352B |doi=10.1038/ncomms8352 |issn=2041-1723 |pmc=4468865 |pmid=26056764}}</ref> Suggestions that these structures represent bacterial ]s<ref name="kaye2008">{{cite journal |last1=Kaye |first1=T.G. |last2=Gaugler |first2=G. |last3=Sawlowicz |first3=Z. |year=2008 |title=Dinosaurian Soft Tissues Interpreted as Bacterial Biofilms |journal=PLOS ONE |volume=3 |issue=7 |page=e2808 |doi=10.1371/journal.pone.0002808|pmid=18665236 |pmc=2483347 |bibcode=2008PLoSO...3.2808K |doi-access=free }}</ref> have been rejected,<ref name="peterson2010">{{cite journal |last1=Peterson |first1=J.E. |last2=Lenczewski |first2=M.E. |last3=Scherer |first3=R.P. |year=2010 |title=Influence of Microbial Biofilms on the Preservation of Primary Soft Tissue in Fossil and Extant Archosaurs |journal=PLOS ONE |volume=5 |issue=10 |page=e13334 |bibcode=2010PLoSO...513334P |doi=10.1371/journal.pone.0013334 |doi-access=free |issn=1932-6203 |pmc=2953520 |pmid=20967227}}</ref> but cross-contamination remains a possibility that is difficult to detect.<ref name="buckley2017">{{cite journal |last1=Buckley |first1=M. |last2=Warwood |first2=S. |last3=van Dongen |first3=B. |last4=Kitchener |first4=A.C. |last5=Manning |first5=P.L. |year=2017 |title=A fossil protein chimera; difficulties in discriminating dinosaur peptide sequences from modern cross-contamination |journal=Proceedings of the Royal Society B |volume=284 |issue=1855 |doi=10.1098/rspb.2017.0544|pmid=28566488 |pmc=5454271 }}</ref>
Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including ]ized ]s, ], ]s, ]s, ]s, impressions of ], ] and ]s.<ref name="softtissue">Dal Sasso, C. and Signore, M. (1998). Exceptional soft-tissue preservation in a theropod dinosaur from Italy. ''Nature'' 292:383-387. </ref><ref>Schweitzer, M.H., Wittmeyer, J.L. and Horner, J.R. (2005). Soft-Tissue Vessels and Cellular Preservation in Tyrannosaurus rex. ''Science'' 307:1952 - 1955. </ref> Many fields of study contribute to our understanding of dinosaurs, including ], ], ], and the ] (of which ] is a sub-discipline).


==Evolutionary history==
Dinosaur remains have been found on every continent on Earth, including ]. Numerous fossils of the same dinosaur species have been found on completely different continents, corroborating the generally-accepted theory that all land masses were at one time connected in a super-continent called ]. Pangaea began to break apart during the ] period roughly 230 million years ago.<ref>Evans, J. (1998). ''Ultimate Visual Dictionary - 1998 Edition''. Dorling Kindersley Books. 66-69. ISBN 1-871854-00-8.</ref>
===Origins and early evolution===
]'' (large), '']'' (small) and a '']'' skull, from the ]]]
Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic epochs, roughly 20&nbsp;million years after the devastating ] wiped out an estimated 96% of all marine species and 70% of terrestrial vertebrate species approximately 252&nbsp;million years ago.<ref name=KPA/><ref name=TannerLucas/> The oldest dinosaur fossils known from substantial remains date to the ] epoch of the Triassic period and have been found primarily in the ] and ]s of Argentina and Brazil, and the ] of Zimbabwe.<ref name="griffin2022">{{cite journal |last1=Griffin |first1=C.T. |last2=Wynd |first2=B.M. |last3=Munyikwa |first3=D. |last4=Broderick |first4=T.J. |last5=Zondo |first5=M. |last6=Tolan |first6=S. |last7=Langer |first7=M.C. |last8=Nesbitt |first8=S.J. |last9=Taruvinga |first9=H.R. |year=2022 |title=Africa's oldest dinosaurs reveal early suppression of dinosaur distribution |journal=Nature |volume=609 |issue=7926 |pages=313–319 |doi=10.1038/s41586-022-05133-x |pmid=36045297 |bibcode=2022Natur.609..313G |s2cid=251977824 |issn=0028-0836}}</ref>


The Ischigualasto Formation (] at 231–230 million years old<ref name="desojo2020">{{cite journal |last1=Desojo |first1=J.B. |last2=Fiorelli |first2=L.E. |last3=Ezcurra |first3=M.D. |last4=Martinelli |first4=A.G. |last5=Ramezani |first5=J. |last6=Da Rosa |first6=A.A.S. |last7=Belén von Baczko |first7=M. |last8=Jimena Trotteyn |first8=M. |last9=Montefeltro |first9=F.C. |last10=Ezpeleta |first10=M. |last11=Langer |first11=M.C. |year=2020 |title=The Late Triassic Ischigualasto Formation at Cerro Las Lajas (La Rioja, Argentina): fossil tetrapods, high-resolution chronostratigraphy, and faunal correlations |journal=Scientific Reports |volume=10 |issue=1 |pages=12782 |doi=10.1038/s41598-020-67854-1 |pmid=32728077 |pmc=7391656 |bibcode=2020NatSR..1012782D}}</ref>) has produced the early saurischian '']'', originally considered a member of the ]<ref name="OARM2010">{{cite journal |last1=Alcober |first1=Oscar A. |last2=Martinez |first2=Ricardo N. |year=2010 |title=A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina |journal=] |location=] |publisher=] |issue=63 |pages=55–81 |doi=10.3897/zookeys.63.550 |pmc=3088398 |issn=1313-2989 |pmid=21594020|bibcode=2010ZooK...63...55A |doi-access=free}}</ref> but now considered to be an early sauropodomorph, along with the herrerasaurids ''Herrerasaurus'' and '']'', and the sauropodomorphs '']'', '']'', and '']''.<ref name="novas2021">{{cite journal |last1=Novas |first1=F.E. |last2=Agnolin |first2=F.L. |last3=Ezcurra |first3=M.D. |last4=Müller |first4=R.T. |last5=Martinelli |first5=A. |last6=Langer |first6=M. |year=2021 |title=Review of the fossil record of early dinosaurs from South America, and its phylogenetic implications |journal=Journal of South American Earth Sciences |volume=110 |pages=103341 |doi=10.1016/j.jsames.2021.103341 |issn=0895-9811 |bibcode=2021JSAES.11003341N}}</ref> ''Eoraptor''{{'s}} likely resemblance to the ] of all dinosaurs suggests that the first dinosaurs would have been small, bipedal ].<ref name="Daemonosaurus">{{cite journal |last1=Nesbitt |first1=Sterling J |last2=Sues |first2=Hans-Dieter |title=The osteology of the early-diverging dinosaur ''Daemonosaurus chauliodus'' (Archosauria: Dinosauria) from the Coelophysis Quarry (Triassic: Rhaetian) of New Mexico and its relationships to other early dinosaurs |journal=Zoological Journal of the Linnean Society |date=2021 |volume=191 |issue=1 |pages=150–179 |doi=10.1093/zoolinnean/zlaa080|doi-access=free }}</ref><ref name=Sereno1999/><ref name=SFRM93/> The Santa Maria Formation (radiometrically dated to be older, at 233.23&nbsp;million years old<ref name=langer18>{{cite journal |last1=Langer |first1=Max C.|last2=Ramezani |first2=Jahandar |last3=Da Rosa |first3=Átila A.S. |title=U-Pb age constraints on dinosaur rise from south Brazil |date=May 2018 |journal=] |location=Amsterdam |publisher=Elsevier |volume=57 |pages=133–140 |doi=10.1016/j.gr.2018.01.005 |bibcode=2018GondR..57..133L |issn=1342-937X}}</ref>) has produced the herrerasaurids '']'' and '']'', along with the sauropodomorphs '']'', '']'', '']'', '']'', '']'', '']'', ''Saturnalia'', and '']''.<ref name="novas2021"/> The Pebbly Arkose Formation, which is of uncertain age but was likely comparable to the other two, has produced the sauropodomorph '']'', along with an unnamed herrerasaurid.<ref name="griffin2022"/>
===The current "dinosaur renaissance"===
{{main|Dinosaur renaissance}}
The field of dinosaur research has enjoyed a surge in activity that began in the 1970s and is ongoing. This was triggered, in part, by ]'s discovery of '']'', an active, vicious ] that may have been ], in marked contrast to the prevailing image of dinosaurs as sluggish and ]. ], arguably the primary scientific discipline involved in dinosaur research, has become a global ]. Major new dinosaur discoveries have been made by paleontologists working in previously unexploited regions, including ], ], ], ], and most significantly in ] (the amazingly well-preserved ] in ] have further solidified the link between dinosaurs and their conjectured living descendants, modern ]s). The widespread application of ], which rigorously analyzes the relationships between biological organisms, has also proved tremendously useful in ] dinosaurs. Cladistic analysis, among other modern techniques, helps to compensate for an often incomplete and fragmentary ].


Less well-preserved remains of the sauropodomorphs '']'' and '']'', along with the early saurischian '']'', are known from the ] and ]s of India.<ref name="novas2011">{{cite journal |last1=Novas |first1=F.E. |last2=Ezcurra |first2=M.D. |last3=Chatterjee |first3=S. |last4=Kutty |first4=T.S. |year=2011 |title=New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |volume=101 |issue=3–4 |pages=333–349 |doi=10.1017/S1755691011020093|bibcode=2010EESTR.101..333N |s2cid=128620874 }}</ref> The Carnian-aged ] of Argentina preserves primitive, dinosaur-like ornithodirans such as '']'' and '']'' in ], making it another important site for understanding dinosaur evolution. These ornithodirans support the model of early dinosaurs as small, bipedal predators.<ref name="novas2021"/><ref name="mariscano2016">{{cite journal |last1=Marsicano |first1=C.A. |last2=Irmis |first2=R.B. |last3=Mancuso |first3=A.C. |last4=Mundil |first4=R. |last5=Chemale |first5=F. |year=2016 |title=The precise temporal calibration of dinosaur origins |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=3 |pages=509–513 |doi=10.1073/pnas.1512541112|pmid=26644579 |pmc=4725541 |bibcode=2016PNAS..113..509M |doi-access=free }}</ref> Dinosaurs may have appeared as early as the ] epoch of the Triassic, approximately 243&nbsp;million years ago, which is the age of '']'' from the ] of Tanzania. However, its known fossils are too fragmentary to identify it as a dinosaur or only a close relative.<ref name=nyasasaurus>{{cite journal |last1=Nesbitt |first1=Sterling J. |last2=Barrett |first2=Paul M. |last3=Werning |first3=Sarah |last4=Sidor |first4=Christian A. |author-link4=Christian Sidor |last5=Charig |first5=Alan J. |author-link5=Alan J. Charig |display-authors=3 |year=2012 |title=The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania |journal=] |volume=9 |issue=1 |page=20120949 |location=London |publisher=Royal Society |doi=10.1098/rsbl.2012.0949 |issn=1744-9561 |pmc=3565515 |pmid=23221875}}</ref> The referral of the Manda Formation to the Anisian is also uncertain. Regardless, dinosaurs existed alongside non-dinosaurian ornithodirans for a period of time, with estimates ranging from 5–10 million years<ref name="marsicano2015">{{cite journal |last1=Marsicano |first1=C.A. |last2=Irmis |first2=R.B. |last3=Mancuso |first3=A.C. |last4=Mundil |first4=R. |last5=Chemale |first5=F. |year=2015 |title=The precise temporal calibration of dinosaur origins |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=3 |pages=509–513 |doi=10.1073/pnas.1512541112 |pmid=26644579 |pmc=4725541 |bibcode=2016PNAS..113..509M |issn=0027-8424 |doi-access=free}}</ref> to 21 million years.<ref name="langer18"/>
{{seealso|List of U.S. state dinosaurs}}


When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of ] and ]s, like ]s and ]s. Their main competitors were the ], such as ]s, ] and rauisuchians, which were more successful than the dinosaurs.<ref>{{cite journal |last1=Brusatte |first1=Stephen L. |author-link1=Stephen L. Brusatte |last2=Benton |first2=Michael J. |last3=Ruta |first3=Marcello |author-link3=Marcello Ruta |last4=Lloyd |first4=Graeme T. |year=2008 |title=Superiority, Competition, and Opportunism in the Evolutionary Radiation of Dinosaurs |url=https://www.pure.ed.ac.uk/ws/files/8232088/PDF_Brusatteetal2008SuperiorityCompetition.pdf |archive-url=https://web.archive.org/web/20180719005836/https://www.pure.ed.ac.uk/ws/files/8232088/PDF_Brusatteetal2008SuperiorityCompetition.pdf |archive-date=2018-07-19 |url-status=live |journal=Science |location=Washington, D.C. |publisher=American Association for the Advancement of Science |volume=321 |issue=5895 |pages=1485–1488 |doi=10.1126/science.1161833 |bibcode=2008Sci...321.1485B |issn=0036-8075 |pmid=18787166 |access-date=October 22, 2019|hdl=20.500.11820/00556baf-6575-44d9-af39-bdd0b072ad2b |s2cid=13393888 }}</ref> Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215&nbsp;million years ago, a variety of ] archosauromorphs, including the ], became extinct. This was followed by the Triassic–Jurassic extinction event (about 201&nbsp;million years ago), that saw the end of most of the other groups of early archosaurs, like aetosaurs, ornithosuchids, ]s, and rauisuchians. Rhynchosaurs and ]s survived (at least in some areas) at least as late as early –mid ] and late Norian or earliest ] ]s, respectively,<ref>{{harvnb|Tanner|Spielmann|Lucas|2013|pp=|loc="The first Norian (Revueltian) rhynchosaur: Bull Canyon Formation, New Mexico, U.S.A." by Justin A. Spielmann, Spencer G. Lucas and Adrian P. Hunt.}}</ref><ref>{{cite journal |last1=Sulej |first1=Tomasz |last2=Niedźwiedzki |first2=Grzegorz |year=2019 |title=An elephant-sized Late Triassic synapsid with erect limbs |journal=Science |location=Washington, D.C. |publisher=American Association for the Advancement of Science |volume=363 |issue=6422 |pages=78–80 |doi=10.1126/science.aal4853 |issn=0036-8075 |pmid=30467179|bibcode=2019Sci...363...78S |s2cid=53716186 |doi-access=free }}</ref> and the exact date of their ] is uncertain. These losses left behind a land fauna of ], dinosaurs, mammals, pterosaurians, and ]s.<ref name=MJB04dino/> The first few lines of early dinosaurs ] through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups that became extinct.<ref name="Letal05"/> Also notably, there was a heightened rate of extinction during the ].<ref>{{cite news |author=<!--Staff writer(s); no by-line.--> |date=April 19, 2018 |title=Fossil tracks in the Alps help explain dinosaur evolution |url=https://www.economist.com/science-and-technology/2018/04/19/fossil-tracks-in-the-alps-help-explain-dinosaur-evolution |url-access=registration |department=Science and Technology |newspaper=] |location=London |issn=0013-0613 |access-date=May 24, 2018}}</ref>
===Classification===
''Main article'': ]


===Evolution and paleobiogeography===
Dinosaurs (including birds) are ]s, like modern ]s. Archosaurs' ] skulls have two holes located where the jaw muscles attach, called ]. Most reptiles (including birds) are diapsids; mammals, with only one temporal fenestra, are called ]s; and ]s, with no temporal fenestra, are ]s. Anatomically, dinosaurs share many other archosaur characteristics, including teeth that grow from sockets rather than as direct extensions of the jawbones. Within the archosaur group, dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodylians sprawl out to either side. All dinosaurs were land animals.
] in the early ] (around 200 million years ago)]]
Dinosaur evolution after the Triassic followed changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass ], and there was a worldwide dinosaur fauna mostly composed of ] carnivores and early sauropodomorph herbivores.<ref name=HCL04/> ] plants (particularly ]s), a potential food source, ] in the Late Triassic. Early sauropodomorphs did not have sophisticated mechanisms for processing food in the mouth, and so must have employed other means of breaking down food farther along the digestive tract.<ref name=FS04/> The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ]ns, ], and ], and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the ] of ] and ] of Tanzania. Dinosaurs in China show some differences, with specialized ] theropods and unusual, long-necked sauropods like '']''.<ref name=HCL04/> Ankylosaurians and ornithopods were also becoming more common, but primitive sauropodomorphs had become extinct. Conifers and ]s were the most common plants. Sauropods, like earlier sauropodomorphs, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential ]-like organs to keep food in the mouth, and jaw motions to grind food.<ref name=FS04/> Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.<ref name=KP04/>


By the ] and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, ]ns, and ] through Europe, North America, and northern ]. These were later supplemented or replaced in Africa by large spinosaurid and ] theropods, and ] and ]n sauropods, also found in ]. In ], maniraptoran coelurosaurians like dromaeosaurids, ], and ]ns became the common theropods, and ] and early ceratopsians like ''Psittacosaurus'' became important herbivores. Meanwhile, ] was home to a fauna of basal ankylosaurians, ]s, and iguanodontians.<ref name=HCL04/> The stegosaurians appear to have gone extinct at some point in the late Early Cretaceous or early ]. A major change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of ]s. At the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of grinding with ], taken to its extreme in hadrosaurids.<ref name=FS04/> Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid '']''.<ref name=serenoetal07/>
Many other types of reptiles lived at the same time as the dinosaurs. Some of these are commonly, but incorrectly, thought of as dinosaurs, including ]s (which are not closely related to the dinosaurs) and ], which developed separately from reptilian ancestors in the late Triassic period.


There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosaurids and various types of smaller maniraptoran theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosaurians. In the southern continents that had made up the now-splitting supercontinent ], ] were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, ] iguanodontians, ] ankylosaurians, and titanosaurian sauropods were prevalent.<ref name=HCL04/> Flowering plants were greatly radiating,<ref name=FS04/> with the first grasses appearing by the end of the Cretaceous.<ref name=PSAS05/> Grinding hadrosaurids and shearing ceratopsians became very diverse across North America and Asia. Theropods were also radiating as herbivores or ]s, with ]ians and ]ns becoming common.<ref name=FS04/>
Collectively, dinosaurs are usually regarded as a ] or an unranked ]. They are divided into two ], the '']'' and the '']'', on the basis of their hip structure. Saurischians ('lizard-hipped', from the ] ''sauros'' (''σαυρος'') meaning 'lizard' and ''ischion'' (''ισχιον'') meaning 'hip joint') are dinosaurs that originally retained the hip structure of their ancestors. They include all the ] (bipedal ]s) and ]s (long-necked ]s). Ornithischians ('bird-hipped', from the ] ''ornitheos'' (''ορνιθειος'') meaning 'of a bird' and ''ischion'' (''ισχιον'') meaning 'hip joint') is the other dinosaurian order, most of which were ]al herbivores. ('''NB:''' the terms "lizard hip" and "bird-hip" are misnomers &mdash; birds evolved from dinosaurs with "lizard hips".)


The Cretaceous–Paleogene extinction event, which occurred approximately 66&nbsp;million years ago at the end of the Cretaceous, caused the extinction of all dinosaur groups except for the neornithine birds. Some other diapsid groups, including ]ns, ], ]ns, turtles, ]s, ]s, ]ns, and ]ns, also survived the event.<ref name=AF04/>
<center><gallery>
Image:Saurischia.png|]n pelvis structure (left side)
Image:Tyrannosaurus pelvis left.JPG|'']'' pelvis (showing saurischian structure - left side)
Image:Ornithischia.png|]n pelvis structure (left side).
Image:Edmontosaurus pelvis left.JPG|'']'' pelvis (showing ornithischian structure - left side)
</gallery></center>


The surviving lineages of neornithine birds, including the ancestors of modern ]s, ], and a variety of ], diversified rapidly at the beginning of the ] period, entering ]s left vacant by the extinction of Mesozoic dinosaur groups such as the arboreal ]s, aquatic ]s, and even the larger terrestrial theropods (in the form of '']'', ], ], ratites, ], ]s, and "]s"). It is often stated that mammals out-competed the neornithines for dominance of most terrestrial niches but many of these groups co-existed with rich mammalian faunas for most of the ] Era.<ref name=lindow>{{harvnb|Dyke|Kaiser|2011|loc=chpt. 14: "Bird Evolution Across the K–Pg Boundary and the Basal Neornithine Diversification" by Bent E. K. Lindow. {{doi|10.1002/9781119990475.ch14}}}}</ref> Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals,<ref name="Cracraft">{{cite journal |last=Cracraft |first=Joel |author-link=Joel Cracraft |year=1968 |title=A Review of the Bathornithidae (Aves, Gruiformes), with Remarks on the Relationships of the Suborder Cariamae |url=https://digitallibrary.amnh.org/bitstream/handle/2246/2536//v2/dspace/ingest/pdfSource/nov/N2326.pdf?sequence=1&isAllowed=y |journal=] |location=New York |publisher=American Museum of Natural History |issue=2326 |pages=1–46 |issn=0003-0082 |hdl=2246/2536 |access-date=October 22, 2019}}</ref><ref>{{cite journal |last1=Alvarenga |first1=Herculano |author1-link=Herculano Marcos Ferraz de Alvarenga |last2=Jones |first2=Washington W. |last3=Rinderknecht |first3=Andrés |date=May 2010 |title=The youngest record of phorusrhacid birds (Aves, Phorusrhacidae) from the late Pleistocene of Uruguay |url=https://www.researchgate.net/publication/233512584 |journal=] |location=] |publisher=] |volume=256 |issue=2 |pages=229–234 |doi=10.1127/0077-7749/2010/0052 |issn=0077-7749 |access-date=October 22, 2019}}</ref> and ratites are still fairly successful as midsized herbivores; eogruiids similarly lasted from the ] to ], becoming extinct only very recently after over 20&nbsp;million years of co-existence with many mammal groups.<ref>{{harvnb|Mayr|2009}}</ref>
The following is a simplified classification of dinosaur families. A more detailed version can be found at ].


==Classification==
The dagger (†) is used to indicate taxa that are ].
{{Main|Dinosaur classification}}


{{multiple image
===Order ]===
|align = right
* †Infraorder ]
|perrow=2/2
* Suborder ]
|total_width = 350
** †Superfamily ]
** †Infraorder ]
*** †Family ]
** (unranked) ]
*** †Superfamily ]
*** †Infraorder ]
*** Infraorder ]
**** †Family ]
**** †Superfamily ]
**** †(unranked) ]
**** (unranked) ]
***** †(unranked) Oviraptoriformes
****** †(unranked) ]
****** †(unranked) ]
***** †(unranked) ]
****** †Family ]
****** †Family ]
***** Class ] (birds)
* †Suborder ]
** †'']''
** †Infraorder ]
** †Infraorder ]
*** †Superfamily ]
*** †Superfamily ]
**** †Family ]
**** †(unranked) ]


|image1 = Saurischia pelvis.png
=== Order ] ===
|alt1 =
* †Suborder ]
|caption1 = ]n pelvis structure (left side)
** †Infraorder ]
** †Infraorder ]
* †(unranked) ]
** †Family ]
** †Suborder ]
*** †Infraorder ]
*** †Infraorder ]
**** †Family ]
**** †Family ]
**** †Family ]
** †Suborder ]
*** †Family ]
*** †Infraorder ]
**** †Family ]
**** †Family ]


|image2 = Tyrannosaurus pelvis left.jpg
==Areas of debate==
|alt2 =
===Warm-bloodedness===
|caption2 = '']'' pelvis (showing saurischian structure&nbsp;– left side)
].]]
{{main|Warm-bloodedness of dinosaurs}}
A vigorous debate on the subject of temperature regulation in dinosaurs has been ongoing since the 1960s. Originally, scientists broadly disagreed as to whether dinosaurs were capable of regulating their body temperatures at all. More recently, dinosaur ]y has become the consensus view, and debate has focused on the mechanisms of temperature regulation.


|image3 = Ornithischia pelvis.png
After dinosaurs were discovered, paleontologists first posited that they were ]ic creatures: "terrible ]s" as their name suggests. This supposed cold-bloodedness implied that dinosaurs were relatively slow, sluggish organisms, comparable to modern reptiles, which need external sources of heat in order to regulate their body temperature. Dinosaur ectothermy remained a prevalent view until ], an early proponent of dinosaur endothermy, published an influential paper on the topic in 1968.
|alt3 =
|caption3 = ]n pelvis structure (left side)


|image4 = Edmontosaurus pelvis left.jpg
Modern evidence indicates that dinosaurs thrived in cooler temperate climates, and that at least some dinosaur species must have regulated their body temperature by internal biological means (perhaps aided by the animals' bulk). Evidence of ]ism in dinosaurs includes the discovery of ] and ] (where they would have experienced a cold, dark six-month winter), the discovery of dinosaurs whose feathers may have provided regulatory insulation, and analysis of blood-vessel structures that are typical of endotherms within dinosaur bone. Skeletal structures suggest that theropods and other dinosaurs had active lifestyles better suited to an endothermic cardiovascular system, while sauropods exhibit fewer endothermic characteristics. It is certainly possible that some dinosaurs were endothermic while others were not. Scientific debate over the specifics continues.<ref>Parsons, K.M. (2001). ''Drawing Out Leviathan''. Indiana University Press. 22-48. ISBN 0-253-33937-5.</ref>
|alt4 =
|caption4 = '']'' pelvis (showing ornithischian structure&nbsp;– left side)
}}


Dinosaurs belong to a group known as archosaurs, which also includes modern crocodilians. Within the archosaur group, dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodilians sprawl out to either side.<ref name="B012"/>
Complicating the debate is the fact that warm-bloodedness can emerge based on more than one mechanism. Most discussions of dinosaur endothermy tend to compare them to average birds or mammals, which expend energy to elevate body temperature above that of the environment. Small birds and mammals also possess ], such as ], ], or ]s, which slows down heat loss. However, large mammals, such as elephants, face a different problem due to their relatively small ratio of surface area to volume (]'s principle). This ratio compares the volume of an animal with the area of its skin: as an animal gets bigger, its surface area increases more slowly than its volume. At a certain point, the amount of heat radiated away through the skin drops below the amount of heat produced inside the body, forcing animals to use additional methods to avoid overheating. In the case of elephants, they are hairless, and have large ears which increase their surface area, and have behavioral adaptations as well (such as using the trunk to spray water on themselves and mud wallowing). These behaviors increase cooling through evaporation.


Collectively, dinosaurs as a clade are divided into two primary branches, Saurischia and Ornithischia. Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all ] sharing a more recent common ancestor with ''Triceratops'' than with Saurischia. Anatomically, these two groups can be distinguished most noticeably by their ] structure. Early saurischians—"lizard-hipped", from the ] ''{{lang|grc-Latn|sauros}}'' ({{lang|grc|σαῦρος}}) meaning "lizard" and ''{{lang|grc-Latn|ischion}}'' ({{lang|grc|ἰσχίον}}) meaning "hip joint"—retained the hip structure of their ancestors, with a pubis bone directed ], or forward.<ref name=MJB00/> This basic form was modified by rotating the pubis backward to varying degrees in several groups (''Herrerasaurus'',<ref name=GSP88/> therizinosauroids,<ref name=clarketal2004/> dromaeosaurids,<ref name=MAPM04/> and birds<ref name=KP04/>). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups).<ref name=theropods/><ref name=sauropodomorphs>{{cite journal |last1=Taylor |first1=Michael P. |author-link1=Michael P. Taylor |last2=Wedel |first2=Mathew J. |author-link2=Matt Wedel |year=2013 |title=Why sauropods had long necks; and why giraffes have short necks |journal=] |location=]; London |volume=1 |page=e36 |doi=10.7717/peerj.36 |issn=2167-8359 |pmid=23638372 |pmc=3628838 |doi-access=free }}</ref>
Large dinosaurs would presumably have had to deal with similar issues; their body size would dictate that they lost heat relatively slowly to the surrounding air, and so could have been what are called ], animals that are warmer than their environments through sheer size rather than through special adaptations like those of birds or mammals. However, so far this theory fails to account for the vast number of dog- and goat-sized dinosaur species which made up the bulk of the ecosystem during the Mesozoic period.


By contrast, ornithischians—"bird-hipped", from the Greek ''ornitheios'' (ὀρνίθειος) meaning "of a bird" and ''ischion'' (ἰσχίον) meaning "hip joint"—had a pelvis that superficially resembled a bird's pelvis: the pubic bone was oriented caudally (rear-pointing). Unlike birds, the ornithischian pubis also usually had an additional forward-pointing process. Ornithischia includes a variety of species that were primarily herbivores.
===Feathered dinosaurs and the bird connection===
{{main|Feathered dinosaurs}}
{{main|Dinosaur-bird connection}}


Despite the terms "bird hip" (Ornithischia) and "lizard hip" (Saurischia), birds are not part of Ornithischia. Birds instead belong to Saurischia, the "lizard-hipped" dinosaurs—birds evolved from earlier dinosaurs with "lizard hips".<ref name="B012"/>
Birds and non-avian dinosaurs share many features. Birds share over a hundred distinct anatomical features with ] dinosaurs, which are generally accepted to have been their closest ancient relatives.<ref>Mayr, G., Pohl, B. and Peters, D.S. (2005). A Well-Preserved Archaeopteryx Specimen with Theropod Features. ''Science'' 310:1483-1486..</ref>


===Taxonomy===
'''Feathers'''
The following is a simplified classification of dinosaur groups based on their evolutionary relationships, and those of the main dinosaur groups Theropoda, Sauropodomorpha and Ornithischia, compiled by Justin Tweet.<ref>{{Cite web|author=Justin Tweet|url=https://equatorialminnesota.blogspot.com/p/blog-page_20.html|title=Classification diagrams|website=Equatorial Minnesota|access-date=2022-09-06}}</ref> Further details and other hypotheses of classification may be found on individual articles.
]'' on display at the ].]]


*Dinosauria
'']'', the first good example of a "feathered dinosaur", was discovered in 1861. The initial specimen was found in the ] in southern Germany, which is a '']'', a rare and remarkable geological formation known for its superbly detailed fossils. Archaeopteryx is a ], with features clearly intermediate between those of modern reptiles and birds. Brought to light just two years after Darwin's seminal '']'', its discovery spurred the nascent debate between proponents of ] and ]. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, specimens are commonly mistaken for '']''.{{citation needed}}
]s; far left: '']'', left: '']'', center background: '']'', center foreground: '']'', right: '']'', far right (large) '']''.]]
:*†] ("bird-hipped"; diverse bipedal and quadrupedal herbivores)
::*†] ("true" ornithischians)
:::*†] (small herbivores/omnivores with prominent ])
::::*†] ("cheeked lizards")
:::::*†] (armored dinosaurs; bipeds and quadrupeds)
::::::*†] (heavy, quadrupedal thyreophorans)
:::::::*†] (spikes and plates as primary armor)
::::::::*†] (small stegosaurs with flank osteoderms and tail clubs)
::::::::*†] (large stegosaurs)
:::::::*†] (]s as primary armor)
::::::::*†] (small, southern ankylosaurs with ]-like tails)
::::::::*†] (mostly spiky, club-less ankylosaurs)
::::::::*†] (characterized by flat scutes)
:::::::::*†] (club-tailed ankylosaurids)
:::::*†] ("new ornithischians")
::::::*†] ("fire teeth")
:::::::*†] ("wondrous lizards")
::::::::*†] (burrowers)
::::::::*†] (large thescelosaurids)
:::::::*†] ("horned feet")
::::::::*†] (characterized by a cranial growth)
]: top left – '']'', top right – '']'', bottom left – '']'', bottom right – '']''.]]
:::::::::*†] (bipeds with domed or knobby growth on skulls)
:::::::::*†] (bipeds and quadrupeds; many had neck frills and horns)
::::::::::*†] (small, frill-less basal ceratopsians)
::::::::::*†] ("new ceratopsians")
:::::::::::*†] (little to no frills, hornless, with robust jaws)
:::::::::::*†] (basal ceratopsians with small frills and stubby horns)
::::::::::::*†] (large-horned ceratopsians)
:::::::::::::*†] (large, elaborately ornamented ceratopsians)
::::::::::::::*†] (ceratopsids with enlarged brow horns)
::::::::::::::*†] (ceratopsids mostly characterized by frill and nasal ornamentation)
:::::::::::::::*†] (centrosaurines with enlarged nasal cavities)
:::::::::::::::*†] (centrosaurines with enlarged nasal horns)
:::::::::::::::*†] (mostly had nasal bosses instead of horns)
::::::::*†] (various sizes; bipeds and quadrupeds; evolved a method of chewing using skull flexibility and numerous teeth)
:::::::::*†] (small European neornithischians)
:::::::::*†] ("iguana teeth"; advanced ornithopods)
::::::::::*†] (with distinctive dentition)
:::::::::::*†] (North American rhabdodontomorphs; bipeds and quadrupeds)
:::::::::::*†] (European rhabdodontomorphs)
::::::::::*†] ("true iguanodonts")
:::::::::::*†] (mostly southern ornithopods with mineralized plates along the ribs; may be thescelosaurids)
:::::::::::*†] ('']'' and more advanced ornithopods)
::::::::::::*†] (mid-sized, small headed)
:::::::::::::*†] (early members mid-sized, stocky)
::::::::::::::*†] ("spiked ]")
:::::::::::::::*†] (ancestrally had a thumb spike; large quadrupedal herbivores, with teeth merged into dental batteries)
::::::::::::::::*†] (hadrosaurids and their closest relatives)
:::::::::::::::::*†] ("duck-billed dinosaurs"; often with crests)
::::::::::::::::::*†] (hadrosaurids with solid, small, no crests)
:::::::::::::::::::*†] (short-crested)
:::::::::::::::::::*†] (enlarged, solid nasal crests)
:::::::::::::::::::*†] (small, spike-like crests)
:::::::::::::::::::*†] (flat-headed saurolophines)
::::::::::::::::::*†] (hadrosaurids often with hollow crests)
:::::::::::::::::::*†] (solid-crested)
:::::::::::::::::::*†] (vertical, tube-like crests)
:::::::::::::::::::*†] (long, backwards-arcing crests)
:::::::::::::::::::*†] (usually rounded crests)
:*]
::*†] (early bipedal carnivores)
]n ]: from left to right '']'', '']'', '']'', and '']'']]
::*†] (herbivores with small heads, long necks, and long tails)
:::*†] (primitive, strictly bipedal "prosauropods")
:::*†] (diverse; bipeds and quadrupeds)
::::*†] ("heavy feet")
:::::*†] (long-necked, primitive sauropodomorphs)
:::::*†] (large, primitive sauropodomorphs)
::::::*†] (heavy, bipeds and quadrupeds)
:::::::*†] (very large and heavy; quadrupedal)
::::::::*†] (gigantic yet lacking several weight-saving adaptations)
::::::::*†] ("heavy lizards")
:::::::::*†] ("true sauropods")
::::::::::*†] (often large, widespread sauropods)
::::::::::*†] ("new sauropods"; columnar limbs)
:::::::::::*†] (skulls and tails elongated; teeth typically narrow and pencil-like)
::::::::::::*†] (short-necked, low-browsing diplodocoids often with high backs)
::::::::::::*†] (whip-tailed)
:::::::::::::*†] (small, short-necked diplodocoids with enlarged cervical and dorsal vertebrae)
:::::::::::::*†] (extremely long-necked)
::::::::::::::*†] (robust cervical vertebrae)
::::::::::::::*†] (long, thin necks)
:::::::::::*†] (boxy skulls; spoon- or pencil-shaped teeth)
::::::::::::*†] ("titan lizard forms")
:::::::::::::*†] (long-necked, long-armed macronarians)
:::::::::::::*†] ("porous vertebrae")
::::::::::::::*†] (stocky, mostly Asian)
::::::::::::::*†] (horse-like skulls; restricted to the Southern Hemisphere; may be titanosaurs)
::::::::::::::*†] (diverse; stocky, with wide hips; most common in the Late Cretaceous of southern continents)
::*] (carnivorous)
:::*] ("new theropods")
::::*†] (early theropods; includes '']'' and close relatives)
::::*†"Dilophosaur-grade neotheropods" (larger kink-snouted dinosaurs)
::::*] ("bird snouts")
:::::*†] (generally elaborately horned carnivores that existed from the Jurassic to Cretaceous periods, originally included Coelophysoidea)
::::::*†] (ceratosaurs with large teeth)
::::::*†] (ceratosaurs exemplified by reduced arms and hands)
:::::::*†] (large abelisauroids with short arms and oftentimes elaborate facial ornamentation)
:::::::*†] (diverse, generally light theropods; may include several obscure taxa)
::::::::*†] (bird-like; omnivorous as juveniles but herbivorous as adults)
::::::::*†] (small carnivores)
:::::*] (stiff-tailed dinosaurs)
::::::*†] (early group of large carnivores)
:::::::*†] (small basal megalosauroids endemic to the Americas)
:::::::*†] (large megalosauroids with powerful arms and hands)
:::::::*†] (crocodile-like, semiaquatic carnivores)
::::::*] ("bird theropods")
:::::::*†] (large meat-eating dinosaurs; megalosauroids sometimes included)
::::::::*†] (primitive Asian allosauroids)
::::::::*†] ('']'' and its very closest relatives)
::::::::*†] (robust allosauroids; includes some of the largest purely terrestrial carnivores)
:::::::*] (feathered theropods, with a range of body sizes and niches)
::::::::*†]? (theropods with large hand claws; potentially tyrannosauroids or neovenatorids)
::::::::*†"Nexus of basal coelurosaurs" (used by Tweet to denote well-known taxa with unstable positions at the base of Coelurosauria)
::::::::*] ("tyrant thieves")
:::::::::*†] (mostly large, primitive coelurosaurs)
::::::::::*†] (tyrannosauroids with head crests)
::::::::::*†] ('']'' and close relatives)
:::::::::*] (bird-like dinosaurs)
::::::::::*†] (small-headed, mostly toothless, omnivorous or possible herbivores)
:::::::::::*†] (very ostrich-like dinosaurs)
::::::::::*] (dinosaurs with pennaceous feathers)
] of six ] ]: from left to right '']'', '']'', '']'', '']'', '']'', and '']'']]
:::::::::::*†] (small hunters with reduced forelimbs)
::::::::::::*†] (insectivores with only one enlarged digit)
:::::::::::*†] (tall, long-necked theropods; omnivores and herbivores)
::::::::::::*†] (larger therizinosaurs)
:::::::::::::*†] (sloth-like herbivores, often with enlarged claws)
:::::::::::*†] (omnivorous, beaked dinosaurs)
::::::::::::*†] (bird-like, basal oviraptorosaurs)
::::::::::::*†] (cassowary-like oviraptorosaurs)
:::::::::::::*†] (toothless oviraptorosaurs known from North America and Asia)
:::::::::::::*†] (characterized by two bony projections at the back of the mouth; exclusive to Asia)
:::::::::::*] (avialans and their closest relatives)
::::::::::::*†] (small tree-climbing theropods with membranous wings)
::::::::::::*†] (toe-clawed dinosaurs; may not form a natural group)
:::::::::::::*†] (small, winged theropods or primitive birds)
:::::::::::::*†] (omnivores; enlarged brain cavities)
:::::::::::::*†] ("raptors")
::::::::::::::*†] (characterized by large wings on both the arms and legs; may have been capable of powered flight)
::::::::::::::*†] (hunters with greatly enlarged sickle claws)
:::::::::::::*†] (piscivores; may be dromaeosaurids)
::::::::::::::*†] (duck-like; potentially semiaquatic)
::::::::::::::*†] (long-snouted)
::::::::::::*] (modern birds and extinct relatives)


===Timeline of major groups===
Since the 1990s, a number of additional ] have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Most of these specimens were unearthed in the ] province in northeastern ], which was part of an island continent during the Cretaceous period. Though feathers have been found only in the ] of the ] and a few other places, it is possible that non-avian dinosaurs elsewhere in the world were also feathered. The lack of widespread fossil evidence for feathered non-avian dinosaurs may be due to the fact that delicate features like skin and feathers are not often preserved by ]ization and thus are absent from the fossil record.


Timeline of major dinosaur groups per {{harvcoltxt|Holtz|2007}}.
The feathered dinosaurs discovered so far include '']'', '']'', '']'', '']'', '']'', '']'', '']'', '']'', and '']''. Dinosaur-like birds like '']'', which are anatomically closer to modern avians, have also been discovered. All of these specimens come from the same formation in northern China. The ] family in particular seems to have been heavily feathered, and at least one dromaeosaurid, '']'', may have been capable of flight.


<timeline>
'''Skeleton'''
ImageSize = width:1000px height:auto barincrement:15px
]'' skeleton at the ].]]
PlotArea = left:10px bottom:50px top:10px right:10px
Because feathers are often associated with birds, feathered dinosaurs are often touted as the ] between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent the more important link for ]s. Furthermore, it is increasingly clear that the relationship between birds and dinosaurs, and the evolution of flight, are more complex topics than previously realized. For example, while it was once believed that birds evolved from dinosaurs in one linear progression, some scientists, most notably ], conclude that dinosaurs such as the ]s may have evolved from birds, losing the power of flight while keeping their feathers in a manner similar to the modern ostrich and other ]s.


Period = from:-251 till:0
Comparison of bird and dinosaur skeletons, as well as ], strengthens the case for the link, particularly for a branch of theropods called ]s. Skeletal similarities include the ], ], ] (semi-lunate ]), ] and ], ], ] and ].
TimeAxis = orientation:horizontal
ScaleMajor = unit:year increment:10 start:-250
ScaleMinor = unit:year increment:1 start:-251
TimeAxis = orientation:hor
AlignBars = justify
Legend = orientation:vertical position:bottom columns:1


Colors =
'''Reproductive biology'''
#legends
]'' skull and upper vertebral column, Palais de la Découverte, Paris.]]
id:CAR value:claret
]'' was a typical "armored dinosaur" of the ] superfamily.]]
id:ANK value:rgb(0.4,0.3,0.196)
A discovery of features in a '']'' ] recently provided even more evidence that dinosaurs and birds evolved from a common ancestor and, for the first time, allowed paleontologists to establish the sex of a dinosaur. When laying eggs, female birds grow a special type of bone in their limbs. This ], which is rich in calcium, forms a layer inside the hard outer bone that is used to make eggshells. The presence of endosteally-derived bone tissues lining the interior marrow cavities of portions of the '']'' specimen's hind limb suggested that ''T. rex'' used similar reproductive strategies, and revealed the specimen to be female.
id:HER value:teal
id:HAD value:green
id:OMN value:blue
id:black value:black
id:white value:white
id:mesozoic value:rgb(0.54,0.54,0.258)
id:triassic value:rgb(0.51,0.17,0.57)
id:earlytriassic value:rgb(0.6,0.22,0.61)
id:middletriassic value:rgb(0.73,0.53,0.71)
id:latetriassic value:rgb(0.78,0.65,0.8)
id:jurassic value:rgb(0.2,0.7,0.79)
id:earlyjurassic value:rgb(0,0.69,0.89)
id:middlejurassic value:rgb(0.52,0.81,0.91)
id:latejurassic value:rgb(0.74,0.89,0.97)
id:cretaceous value:rgb(0.5,0.78,0.31)
id:earlycretaceous value:rgb(0.63,0.78,0.65)
id:latecretaceous value:rgb(0.74,0.82,0.37)
id:cenozoic value:rgb(0.54,0.54,0.258)
id:paleogene value:rgb(0.99,0.6,0.32)
id:paleocene value:rgb(0.99,0.65,0.37)
id:eocene value:rgb(0.99,0.71,0.42)
id:oligocene value:rgb(0.99,0.75,0.48)
id:neogene value:rgb(0.999999,0.9,0.1)
id:miocene value:rgb(0.999999,0.999999,0)
id:pliocene value:rgb(0.97,0.98,0.68)
id:quaternary value:rgb(0.98,0.98,0.5)
id:pleistocene value:rgb(0.999999,0.95,0.68)
id:holocene value:rgb(0.999,0.95,0.88)
id:her value:red Legend:Herrerasauria
id:pur value:purple Legend:Sauropodomorpha
id:ther value:orange Legend:Theropoda
id:orn value:green Legend:Ornithischia
Legend = columns:1 left:100 top:20 columnwidth:100


A dinosaur embryo was found without teeth, suggesting that some parental care was required to feed the young dinosaur. It is also possible that the adult dinosaurs regurgitated into a young dinosaur's mouth to provide sustenance, a behavior that is also characteristic of numerous modern bird species.
BarData=
bar:eratop
bar:space
bar:periodtop
bar:space
bar:NAM1
bar:NAM2
bar:NAM3
bar:NAM4
bar:NAM5
bar:NAM6
bar:NAM7
bar:NAM8
bar:NAM9
bar:NAM10
bar:NAM11
bar:NAM12
bar:NAM13
bar:NAM14
bar:NAM15
bar:NAM16
bar:NAM17
bar:NAM18
bar:NAM19
bar:NAM20
bar:NAM21
bar:NAM22
bar:NAM23
bar:NAM24
bar:NAM25
bar:NAM26
bar:NAM27
bar:NAM28
bar:NAM29
bar:NAM30


bar:space
'''Lungs'''
bar:period
bar:space
bar:era


PlotData=
Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to an investigation which was led by ] of ]. The lungs of theropod dinosaurs (carnivores that walked on two legs and had birdlike feet) likely pumped air into hollow sacs in their ]s, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said. The study was funded in part by the ].<ref>O'Connor, P.M. and Claessens, L.P.A.M. (2005). Basic avian pulmonary design and flow-through ventilation in non-avian theropod dinosaurs. ''Nature'' 436:253.</ref>
align:center textcolor:black fontsize:M mark:(line,black) width:25
shift:(2,-5)
bar:periodtop
from: -251 till: -245 color:earlytriassic text:]
from: -245 till: -228 color:middletriassic text:]
from: -228 till: -199.6 color:latetriassic text:]
from: -199.6 till: -175.6 color:earlyjurassic text:]
from: -175.6 till: -161.2 color:middlejurassic text:]
from: -161.2 till: -145.5 color:latejurassic text:]
from: -145.5 till: -99.6 color:earlycretaceous text:]
from: -99.6 till: -65.5 color:latecretaceous text:]
from: -65.5 till: -55.8 color:paleocene text:]
from: -55.8 till: -33.9 color:eocene text:]
from: -33.9 till: -23.03 color:oligocene text:]
from: -23.03 till: -5.332 color:miocene text:]
from: -5.332 till: -2.588 color:pliocene text:]
from: -2.588 till: -0.0117 color:pleistocene text:]
from: -0.0117 till: 0 color:holocene text:]


bar:eratop
'''Heart and sleeping posture'''
from: -251 till: -199.6 color:triassic text:]
from: -199.6 till: -145.5 color:jurassic text:]
from: -145.5 till: -65.5 color:cretaceous text:]
from: -65.5 till: -23.03 color:paleogene text:]
from: -23.03 till: -2.588 color:neogene text:]
from: -2.588 till: 0 color:quaternary text:]


PlotData=
Modern ] (CT) scans of dinosaur chest cavities (conducted in 2000) found the apparent remnants of complex four-chambered hearts, much like those found in today's mammals and birds. A recently discovered ] fossil demonstrates that the dinosaurs slept like certain modern birds, with their heads tucked under their arms.<ref>Xu, X. and Norell, M.A. (2004). A new troodontid dinosaur from China with avian-like sleeping posture. ''Nature'' 431:838-841..</ref> This behavior, which may have helped to keep the head warm, is also characteristic of modern birds.
align:left fontsize:M mark:(line,white) width:5 anchor:till shift:(5,-4)


color:her bar:NAM1 from:-233.23 till:-210 text:]
'''Gizzard'''
color:pur bar:NAM2 from:-231.4 till:-208 text:]
color:pur bar:NAM3 from:-225 till:-190 text:]
color:pur bar:NAM4 from:-228 till:-213 text:]
color:pur bar:NAM5 from:-227 till:-176 text:]
color:pur bar:NAM6 from:-183 till:-175 text:]
color:pur bar:NAM7 from:-168 till:-125 text:]
color:pur bar:NAM8 from:-175 till:-150 text:]
color:pur bar:NAM9 from:-174 till:-93 text:]
color:pur bar:NAM10 from:-157 till:-93 text:]
color:pur bar:NAM11 from:-140 till:-66 text:]
color:ther bar:NAM12 from:-221 till:-183 text:]
color:ther bar:NAM13 from:-199.3 till:-66 text:]
color:ther bar:NAM14 from:-170 till:-85 text:]
color:ther bar:NAM15 from:-175.6 till:-88 text:]
color:ther bar:NAM16 from:-130 till:-66 text:]
color:ther bar:NAM17 from:-166 till:-66 text:]
color:ther bar:NAM18 from:-151.5 till:-108 text:]
color:ther bar:NAM19 from:-140 till:-66 text:]
color:ther bar:NAM20 from:-160 till:-66 text:]
color:ther bar:NAM21 from:-139 till:-66 text:]
color:ther bar:NAM22 from:-130 till:-66 text:]
color:ther bar:NAM23 from:-167 till:-66 text:]
color:ther bar:NAM24 from:-155 till:0 shift:(-45,5) text:]
color:orn bar:NAM25 from:-200 till:-140 text:]
color:orn bar:NAM26 from:-169 till:-100 text:]
color:orn bar:NAM27 from:-170.3 till:-66 text:]
color:orn bar:NAM28 from:-92 till:-66 text:]
color:orn bar:NAM29 from:-161 till:-66 text:]
color:orn bar:NAM30 from:-164 till:-66 text:]


PlotData=
Another piece of evidence that birds and dinosaurs are closely related is the use of ] stones. These stones are swallowed by animals to aid digestion and break down food and hard fibres once they enter the stomach. When found in association with ]s, gizzard stones are called ]s. Because a particular stone could have been swallowed at one location before being carried to another during migration, paleontologists sometimes use the stones found in dinosaur stomachs to establish possible ] routes.
align:center textcolor:black fontsize:M mark:(line,black) width:25
shift:(2,-5)


bar:period
===Evidence for Cenozoic dinosaurs===
from: -251 till: -245 color:earlytriassic text:]
In 2002, paleontologists Zielinski and Budahn reported the discovery of a single ] leg bone fossil in the San Juan Basin, New Mexico. The formation in which the bone was discovered has been dated to the early ] epoch approximately 64.5 million years ago. If the bone was not re-deposited into that ] by weathering action, it would provide evidence that some dinosaur populations may have survived at least a half million years into the Cenozoic Era.<ref>Fassett, J, R.A. Zielinski, & J.R. Budahn. (2002). Dinosaurs that did not die; evidence for Paleocene dinosaurs in the Ojo Alamo Sandstone, San Juan Basin, New Mexico. In: Catastrophic events and mass extinctions; impacts and beyond. (Eds. Koeberl, C. & K. MacLeod): ''Special Paper - Geological Society of America'' 356: 307-336.</ref>
from: -245 till: -228 color:middletriassic text:]
from: -228 till: -199.6 color:latetriassic text:]
from: -199.6 till: -175.6 color:earlyjurassic text:]
from: -175.6 till: -161.2 color:middlejurassic text:]
from: -161.2 till: -145.5 color:latejurassic text:]
from: -145.5 till: -99.6 color:earlycretaceous text:]
from: -99.6 till: -65.5 color:latecretaceous text:]
from: -65.5 till: -55.8 color:paleocene text:]
from: -55.8 till: -33.9 color:eocene text:]
from: -33.9 till: -23.03 color:oligocene text:]
from: -23.03 till: -5.332 color:miocene text:]
from: -5.332 till: -2.588 color:pliocene text:]
from: -2.588 till: -0.0117 color:pleistocene text:]
from: -0.0117 till: 0 color:holocene text:]


bar:era
===Bringing dinosaurs back to life===
from: -251 till: -199.6 color:triassic text:]
] model of '']''.]]
from: -199.6 till: -145.5 color:jurassic text:]
from: -145.5 till: -65.5 color:cretaceous text:]
from: -65.5 till: -23.03 color:paleogene text:]
from: -23.03 till: -2.588 color:neogene text:]
from: -2.588 till: 0 color:quaternary text:]


</timeline>
There has been much speculation about the use of technology to bring dinosaurs back to life. In ]'s book '']'' (later adapted into a ]), which popularized the idea, scientists use blood from fossilized ]s that have been suspended in ] since the Mesozoic to reconstruct the ] of dinosaurs, filling chromosomal gaps with modern ] genes. It is probably impossible to resurrect dinosaurs in this manner. One problem with the amber extraction method is that DNA decays over time by exposure to air, water and radiation, making it unlikely that such an approach would recover any useful DNA (DNA decay can be measured by a ] test).


==Paleobiology==
The successful extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions, but upon further inspection and ], neither of these reports could be confirmed.<ref>Wang, H., Yan, Z. and Jin, D. (1997). Reanalysis of published DNA sequence amplified from Cretaceous dinosaur egg fossil. ''Molecular Biology and Evolution''. 14:589-591. .</ref> However, a functional visual ] of a (theoretical) dinosaur has been inferred using analytical phylogenetic reconstruction methods on gene sequences of still-living related species (reptiles and birds).<ref>Chang, B.S.W., Jönsson, K., Kazmi, M.A., Donoghue, M.J. and Sakmar, T.P. (2002). Recreating a Functional Ancestral Archosaur Visual Pigment. ''Molecular Biology and Evolution'' 19:1483-1489. .</ref>
Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, ], ]s, ]s, ]s, impressions of skin, ]s and other ]s.<ref name="softtissue">{{cite journal |last1=Dal Sasso |first1=Cristiano |author-link1=Cristiano Dal Sasso |last2=Signore |first2=Marco |date=March 26, 1998 |title=Exceptional soft-tissue preservation in a theropod dinosaur from Italy |journal=Nature |location=London |publisher=Nature Research |volume=392 |issue=6674 |pages=383–387 |doi=10.1038/32884 |bibcode=1998Natur.392..383D |s2cid=4325093 |issn=0028-0836|url=http://doc.rero.ch/record/14901/files/PAL_E2043.pdf |archive-url=https://web.archive.org/web/20160920170653/https://doc.rero.ch/record/14901/files/PAL_E2043.pdf |archive-date=2016-09-20 |url-status=live }}</ref><ref name="Schweitzer2005"/> Many fields of study contribute to our understanding of dinosaurs, including ] (especially ]), ], ], and the ]s (of which ] is a sub-discipline).<ref name=alexander2006/><ref name=dinobiology>{{cite journal |last1=Farlow |first1=James O. |last2=Dodson |first2=Peter |last3=Chinsamy |first3=Anusuya |author-link3=Anusuya Chinsamy-Turan |date=November 1995 |title=Dinosaur Biology |journal=] |location=] |publisher=] |volume=26 |pages=445–471 |doi=10.1146/annurev.es.26.110195.002305 |issn=1545-2069}}</ref> Two topics of particular interest and study have been dinosaur size and behavior.<ref>{{harvnb|Weishampel|Dodson|Osmólska|2004}}</ref>


===Size===
Even if dinosaur DNA could be reconstructed, it would be exceedingly difficult to "grow" dinosaurs using current technology since no closely related species exist to provide ]s or a suitable environment for ].
{{Main|Dinosaur size}}
]s:{{legend|#ae2032|] ('']'')}}{{legend|#cc48cc|]a ('']'')}}{{legend|#50da50|] ('']'')}}
{{legend|#f8863b|] ('']'')}}
{{legend|#254c87|] ('']'')}}]]


Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic, Late Jurassic and Cretaceous.<ref name=Sereno1999/> Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the {{convert|100|to|1000|kg|lb|abbr=on|adj=on}} category when sorted by estimated weight into categories based on ], whereas ] predatory carnivoran mammals peak in the {{convert|10|to|100|kg|lb|abbr=on|adj=on}} category.<ref name=JF93/> The ] of Mesozoic dinosaur body masses is between {{convert|1|and|10|MT|ST}}.<ref name=Peczkis1994/> This contrasts sharply with the average size of Cenozoic mammals, estimated by the ] as about {{convert|2|to|5|kg|lb|abbr=on}}.<ref name=NMNH/>
===Soft tissue in dinosaur fossils===
One of the best examples of soft tissue impressions in a fossil dinosaur was discovered in ], ]. The discovery was reported in 1998, and described the specimen of a small, very young ], ''] samniticus''. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this immature dinosaur.<ref name="softtissue" />


The sauropods were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest was an order of magnitude more massive than anything else that has since walked the Earth. Giant prehistoric mammals such as '']'' (the largest land mammal ever) were dwarfed by the giant sauropods, and only modern whales approach or surpass them in size.<ref name="sander2011">{{cite journal |last1=Sander |first1=P. Martin |last2=Christian |first2=Andreas |last3=Clauss |first3=Marcus |last4=Fechner |first4=Regina |last5=Gee |first5=Carole T. |last6=Griebeler |first6=Eva-Maria |last7=Gunga |first7=Hanns-Christian |last8=Hummel |first8=Jürgen |last9=Mallison |first9=Heinrich |display-authors=3 |date=February 2011 |title=Biology of the sauropod dinosaurs: the evolution of gigantism |journal=Biological Reviews |location=Cambridge |publisher=Cambridge Philosophical Society |volume=86 |issue=1 |pages=117–155 |doi=10.1111/j.1469-185X.2010.00137.x |issn=1464-7931 |pmid=21251189 |pmc=3045712}}</ref> There are several proposed advantages for the large size of sauropods, including protection from predation, reduction of energy use, and longevity, but it may be that the most important advantage was dietary. Large animals are more efficient at digestion than small animals, because food spends more time in their digestive systems. This also permits them to subsist on food with lower nutritive value than smaller animals. Sauropod remains are mostly found in rock formations interpreted as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse would have been advantageous in such environments.<ref name=KC06/>
In the March 2005 issue of '']'', Dr. Mary Higby Schweitzer and her team announced the discovery of flexible material resembling actual soft tissue inside a 68-million-year-old '']'' leg ] from the ] in ]. After recovery, the tissue was rehydrated by the science team.


====Largest and smallest====
When the fossilized bone was treated over several weeks to remove mineral content from the fossilized bone marrow cavity (a process called demineralization), Schweitzer found evidence of intact structures such as ]s, bone matrix, and connective tissue (bone fibers). Scrutiny under the microscope further revealed that the putative dinosaur soft tissue had retained fine structures (microstructures) even at the cellular level. The exact nature and composition of this material are not yet clear, although many news reports immediately linked it with the movie '']''. Interpretation of the artifact is ongoing, and the relative importance of Dr. Schweitzer's discovery is not yet clear.<ref>Schweitzer, M.H., Wittmeyer, J.L. and Horner, J.R. (2005). Soft-Tissue Vessels and Cellular Preservation in Tyrannosaurus rex. ''Science'' 307:1952-1955. Also covers the ]. </ref>
Scientists will probably never be certain of the ] to have ever existed. This is because only a tiny percentage of animals were ever fossilized and most of these remain buried in the earth. Few non-avian dinosaur specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork.<ref>{{harvnb|Paul|2010}}</ref>


]'' to the average human]]
==Extinction theories==
{{main|Cretaceous-Tertiary extinction event}}


The tallest and heaviest dinosaur known from good skeletons is '']'' (previously classified as a species of '']''). Its remains were discovered in Tanzania between 1907 and 1912. Bones from several similar-sized individuals were incorporated into the skeleton now mounted and on display at the ] in ];<ref name=EC68/> this mount is {{convert|12|m|ft|sp=us}} tall and {{convert|21.8|to|22.5|m|ft|sp=us}} long,<ref>{{cite journal |last1=Mazzetta |first1=Gerardo V. |last2=Christiansenb |first2=Per |last3=Fariñaa |first3=Richard A. |year=2004 |title=Giants and Bizarres: Body Size of Some Southern South American Cretaceous Dinosaurs |url=http://www.miketaylor.org.uk/tmp/papers/Mazzetta-et-al_04_SA-dino-body-size.pdf |archive-url=https://web.archive.org/web/20090225155217/http://www.miketaylor.org.uk/tmp/papers/Mazzetta-et-al_04_SA-dino-body-size.pdf |archive-date=2009-02-25 |url-status=live |journal=Historical Biology |location=Milton Park, Oxfordshire |publisher=Taylor & Francis |volume=16 |issue=2–4 |pages=71–83 |doi=10.1080/08912960410001715132 |bibcode=2004HBio...16...71M |citeseerx=10.1.1.694.1650 |s2cid=56028251 |issn=0891-2963}}</ref><ref>{{cite journal |last=Janensch |first=Werner |author-link=Werner Janensch |year=1950 |others=Translation by Gerhard Maier |title=Die Skelettrekonstruktion von ''Brachiosaurus brancai'' |trans-title=The Skeleton Reconstruction of Brachiosaurus brancai |url=https://paleoglot.org/files/Janensch1950b.pdf |url-status=live |journal=Palaeontographica |location=Stuttgart |publisher=] |volume=Suplement VII |issue=1. Reihe, Teil 3, Lieferung 2 |pages=97–103 |oclc=45923346 |archive-url=https://web.archive.org/web/20170711052046/https://paleoglot.org/files/Janensch1950b.pdf |archive-date=July 11, 2017 |access-date=October 24, 2019}}</ref> and would have belonged to an animal that weighed between {{gaps|30|000}} and {{gaps|60|000}}&nbsp;kilograms ({{gaps|70|000}} and {{gaps|130|000}}&nbsp;lb). The longest complete dinosaur is the {{convert|27|m|ft|sp=us}} long ''Diplodocus'', which was discovered in ] in the ] and displayed in ]'s ] in 1907.<ref name=lucas04>{{cite conference |last1=Lucas |first1=Spencer G. |last2=Herne |first2=Matthew C. |last3=Hecket |first3=Andrew B. |last4=Hunt |first4=Adrian P. |last5=Sullivan |first5=Robert M. |display-authors=3 |year=2004 |title=Reappraisal of ''Seismosaurus'', a Late Jurassic Sauropod Dinosaur From New Mexico |url=https://gsa.confex.com/gsa/2004AM/finalprogram/abstract_77727.htm |url-status=live |conference=2004 Denver Annual Meeting (November 7–10, 2004) |conference-url=https://gsa.confex.com/gsa/2004AM/webprogram/start.html |volume=36 |publisher=Geological Society of America |location=Boulder, CO |page=422 |id=Paper No. 181-4 |oclc=62334058 |archive-url=https://web.archive.org/web/20191008110318/https://gsa.confex.com/gsa/2004AM/finalprogram/abstract_77727.htm |archive-date=October 8, 2019 |access-date=October 25, 2019}}</ref> The longest dinosaur known from good fossil material is '']'': the skeleton mount in the American Museum of Natural History in ] is {{convert|37|meters|feet}} long. The ] in ], Argentina, has an '']'' reconstructed skeleton mount that is {{convert|39.7|m|feet|sp=us}} long.<ref name="PLOS One">{{cite journal |last1=Sellers |first1=William Irvin. |last2=Margetts |first2=Lee |last3=Coria |first3=Rodolfo Aníbal |author3-link=Rodolfo Coria |last4=Manning |first4=Phillip Lars |year=2013 |editor1-last=Carrier |editor1-first=David |title=March of the Titans: The Locomotor Capabilities of Sauropod Dinosaurs |doi=10.1371/journal.pone.0078733 |journal=PLOS ONE |location=San Francisco, CA |publisher=PLOS |volume=8 |issue=10 |page=e78733 |pmid=24348896 |pmc=3864407|bibcode=2013PLoSO...878733S |issn=1932-6203|doi-access=free }}</ref>
The sudden ] of the non-avian dinosaurs, which occurred around 65 million years ago, is one of the most intriguing mysteries in ]. Many other groups of animals also became extinct at this time, including ]s (]-like ]s), ]s, plesiosaurs, pterosaurs, herbivorous ]s and ]s, most birds, and many groups of mammals.<ref name="changes">(Nov 2000). ''Earthwatch'' :6-13.</ref> The nature of the event that caused this mass extinction has been extensively studied since the 1970s. At present, several related theories are broadly supported by paleontologists.
]
]'', potentially the largest terrestrial animal to ever exist.]]
There were larger dinosaurs, but knowledge of them is based entirely on a small number of fragmentary fossils. Most of the largest herbivorous specimens on record were discovered in the 1970s or later, and include the massive ''Argentinosaurus'', which may have weighed {{convert|80000|to|100000|kg|ST|sp=us|abbr=off|comma=gaps}} and reached lengths of {{convert|30|to|40|m|ft|sp=us}}; some of the longest were the {{convert|33.5|m|ft|sp=us|adj=mid}} long ''Diplodocus hallorum''<ref name=KC06/> (formerly ''Seismosaurus''), the {{convert|33|to|34|m|ft|sp=us|adj=mid}} long '']'',<ref name=LHW07/> and {{convert|37|m|feet|sp=us|adj=mid}} long ''Patagotitan''; and the tallest, the {{convert|18|m|ft|sp=us|adj=mid}} tall '']'', which could have reached a sixth-floor window. There were a few dinosaurs that was considered either the heaviest and longest. The most famous one include '']'', known only from a now lost partial vertebral ] described in 1878. Extrapolating from the illustration of this bone, the animal may have been {{convert|58|m|ft|sp=us}} long and weighed {{cvt|122400|kg|lb|comma=gaps}}.<ref name=KC06/> However, recent research have placed ''Amphicoelias'' from the long, gracile diplodocid to the shorter but much stockier rebbachisaurid. Now renamed as '']'', this sauropod now stands as much as {{convert|40|m|ft|sp=us}} long and weigh as much as {{cvt|120000|kg|lb|comma=gaps}}.<ref name="carpenter2018">{{cite journal | title=Maraapunisaurus fragillimus, N.G. (formerly Amphicoelias fragillimus), a basal Rebbachisaurid from the Morrison Formation (Upper Jurassic) of Colorado | author=Carpenter, Kenneth | journal=Geology of the Intermountain West | year=2018 | volume=5 | pages=227–244 | doi=10.31711/giw.v5i0.28 | doi-access=free }}</ref><ref>{{Cite journal |last=Paul|first=Gregory S.|date=2019|title=Determining the largest known land animal: A critical comparison of differing methods for restoring the volume and mass of extinct animals |url=http://www.gspauldino.com/Titanomass.pdf |journal=Annals of the Carnegie Museum |volume=85 |issue=4|pages=335–358|doi=10.2992/007.085.0403|s2cid=210840060}}</ref> Another contender of this title includes '']'', a controversial taxon that was recently confirmed to exist after archived photos were uncovered.<ref>{{Cite journal |last1=Pal |first1=Saurabh |last2=Ayyasami |first2=Krishnan |date=27 June 2022 |title=The lost titan of Cauvery |journal=] |language=en |volume=38 |issue=3 |pages=112–116 |doi=10.1111/gto.12390 |bibcode=2022GeolT..38..112P |s2cid=250056201 |issn=0266-6979}}</ref> ''Bruhathkayosaurus'' was a titanosaur and would have most likely weighed more than even ''Marrapunisaurus''. Recent size estimates in 2023 have placed this sauropod reaching lengths of up to {{cvt|44|m|ft}} long and a colossal weight range of around {{cvt|110000–170000|kg|lb|comma=gaps}}, if these upper estimates up true, ''Bruhathkayosaurus'' would have rivaled the '']'' and '']'' as one of the largest animals to have ever existed.<ref name="Bruhathkayosaurus2023">{{Cite journal |last1=Paul |first1=Gregory S. |last2=Larramendi |first2=Asier |date=11 April 2023 |title=Body mass estimate of ''Bruhathkayosaurus'' and other fragmentary sauropod remains suggest the largest land animals were about as big as the greatest whales |journal=Lethaia |language=en |volume=56 |issue=2 |pages=1–11 |doi=10.18261/let.56.2.5 |bibcode=2023Letha..56..2.5P |s2cid=259782734 |issn=0024-1164}}</ref>


The largest carnivorous dinosaur was '']'', reaching a length of {{convert|12.6|to|18|m|ft|sp=us}} and weighing {{convert|7|to|20.9|MT|ST}}.<ref name=SMBM06/><ref name=TH07/> Other large carnivorous theropods included '']'', '']'', and ''Tyrannosaurus''.<ref name=TH07/> '']'' and '']'' were among the tallest of the theropods. The largest ornithischian dinosaur was probably the hadrosaurid '']'' which measured {{convert|16.6|m|feet|sp=us}}.<ref>{{cite journal |last1=Zhao |first1=Xijin |author1-link=Zhao Xijin |last2=Li |first2=Dunjing |last3=Han |first3=Gang |last4=Zhao |first4=Huaxi |last5=Liu |first5=Fengguang |last6=Li |first6=Laijin |last7=Fang |first7=Xiaosi |display-authors=3 |year=2007 |title=Zhuchengosaurus maximus from Shandong Province |journal=Acta Geoscientia Sinica |location=] |publisher=] |volume=28 |issue=2 |pages=111–122 |issn=1006-3021}}</ref> The largest individuals may have weighed as much as {{convert|16|MT|ST}}.<ref>{{harvnb|Weishampel|Dodson|Osmólska|2004|pp=438–463|loc=chpt. 20: "Hadrosauridae" by John R. Horner David B. Weishampel, and Catherine A. Forster.}}</ref>
===Asteroid collision===
] at the tip of the ], the impact of which may have caused the dinosaur extinction.]]
The asteroid collision theory, which was first proposed by ] in the late 1970s, links the ] at the end of the Cretaceous period to a ] impact approximately 65.5 million years ago. Alvarez proposed that a sudden increase in ] levels, recorded around the world in the period's rock stratum, was direct evidence of the impact. The bulk of the evidence now suggests that a 5-15 km wide ] hit in the vicinity of the ], creating the 170&nbsp;km-wide ] and triggering the ]. Scientists are not certain whether dinosaurs were thriving or declining before the impact event. Some scientists propose that the meteorite caused a long and unnatural drop in Earth's atmospheric temperature, while others claim that it would have instead created an unusual heat wave.


], the smallest known dinosaur]]
Although the speed of extinction cannot be deduced from the fossil record alone, various models suggest that the extinction was extremely rapid. The consensus among scientists who support this theory is that the impact caused extinctions both directly (by ] from the meteorite impact) and also indirectly (via a worldwide cooling brought about when matter ejected from the impact crater reflected thermal radiation from the sun).
The smallest dinosaur known is the ],<ref>{{harvnb|Norell|Gaffney|Dingus|2000}}</ref> with a length of only {{convert|5|cm|in|sp=us}} and mass of around {{convert|1.8|g|oz|abbr=on}}.<ref>{{cite web |url=https://www.birds.com/species/a-b/bee-hummingbird/ |url-status=live |title=Bee Hummingbird (''Mellisuga helenae'') |website=Birds.com |publisher=Paley Media |archive-url=https://web.archive.org/web/20150403005328/https://www.birds.com/species/a-b/bee-hummingbird/ |archive-date=April 3, 2015 |access-date=October 27, 2019}}</ref> The smallest known non-] dinosaurs were about the size of ]s and were those theropods most closely related to birds.<ref name=zhang2008/> For example, '']'' is currently the smallest non-avialan dinosaur described from an adult specimen, with an estimated weight of {{convert|110|g|oz|abbr=on}}<ref name=anchiadvance/> and a total skeletal length of {{convert|34|cm|ft|sp=us}}.<ref name=zhang2008/><ref name=anchiadvance/> The smallest herbivorous non-avialan dinosaurs included '']'' and '']'', at about {{convert|60|cm|ft|sp=us}} long each.<ref name=Holtz2007/><ref name="butler&zhao2009"/>


===Behavior===
===Multiple collisions&mdash;the Oort cloud===
]'' was discovered in 1978]]
While similar to Alvarez's impact theory (which involved a single asteroid or comet), this theory proposes that a stream of comets was dislodged from the ] due to the gravitational disruption caused by a passing star. One or more of these objects then collided with the Earth at approximately the same time, causing the worldwide extinction. As with the impact of a single asteroid, the end result of this comet bombardment would have been a sudden drop in global temperatures, followed by a protracted cool period.<ref>Koeberl, C. and MacLeod, K.G. (2002). ''Catastrophic Events and Mass Extinctions''. Geological Society of America. ISBN 0-8137-2356-6.</ref>


Many modern birds are highly social, often found living in flocks. There is general agreement that some behaviors that are common in birds, as well as in ]s (closest living relatives of birds), were also common among extinct dinosaur groups. Interpretations of behavior in fossil species are generally based on the pose of skeletons and their ], ]s of their biomechanics, and comparisons with modern animals in similar ecological niches.<ref name=alexander2006/>
===Environment changes===
At the peak of the dinosaur era, there were no polar ice caps, and sea levels are estimated to have been from 100 to 250 metres (330 to 820 feet) higher than they are today. The planet's temperature was also much more uniform, with only 25 degrees Celsius separating average polar temperatures from those at the equator. On average, atmospheric temperatures were also much warmer; the poles, for example, were 50 °C warmer than today.<ref> The effect climate change may have had on the extinction of the Dinosaurs</ref><ref> Sea levels during the dinosaur era; ]; November 29, 2005</ref>


The first potential evidence for ]ing or ] as a widespread behavior common to many dinosaur groups in addition to birds was the 1878 discovery of 31&nbsp;''Iguanodon'', ornithischians that were then thought to have perished together in ], ], after they fell into a deep, flooded ] and drowned.<ref name=Yans/> Other mass-death sites have been discovered subsequently. Those, along with multiple trackways, suggest that gregarious behavior was common in many early dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that duck-billed (hadrosaurids) may have moved in great herds, like the ] or the African ]. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in ], England,<ref name=Day2002/> although there is no evidence for specific herd structures.<ref name=JLW05/> Congregating into herds may have evolved for defense, for ] purposes, or to provide protection for young. There is evidence that many types of slow-growing dinosaurs, including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. One example is a site in ] that has yielded remains of over 20 '']'', from one to seven years old. This assemblage is interpreted as a social group that was trapped in mud.<ref name=DVetal08sino/> The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as ]s working together to bring down large prey.<ref name=LG93/><ref name="maxwell&ostrom1995"/> However, this lifestyle is uncommon among modern birds, crocodiles, and other reptiles, and the ] evidence suggesting mammal-like pack hunting in such theropods as ''Deinonychus'' and '']'' can also be interpreted as the results of fatal disputes between feeding animals, as is seen in many modern diapsid predators.<ref name=RB07/>
The atmosphere's composition during the dinosaur era was vastly different as well. Carbon dioxide levels were up to 12 times higher than today's levels, and oxygen formed 32 to 35% of the atmosphere, as compared with 21% today. However, by the late ], the environment was changing dramatically. Volcanic activity was decreasing, which led to a cooling trend as levels of atmospheric carbon dioxide dropped. Oxygen levels in the atmosphere also started to fluctuate and would ultimately fall considerably. Some scientists hypothesize that climate change, combined with lower oxygen levels, might have led directly to the demise of many species. If the dinosaurs had respiratory systems similar to those commonly found in modern birds, it may have been particularly difficult for them to cope with reduced respiratory efficiency, given the enormous oxygen demands of their very large bodies.<ref name="changes" />


]'' engaged in ]]]
==History of discovery==
The crests and frills of some dinosaurs, like the ]ns, theropods and ]s, may have been too fragile to be used for active defense, and so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and ]. Head wounds from bites suggest that theropods, at least, engaged in active aggressive confrontations.<ref name=PC98/>
Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese, whose own word for dinosaur is ''konglong'' (恐龍, or "terrible dragon"), considered them to be ] ]s and documented them as such. For example, ''Hua Yang Guo Zhi'', a book written by Zhang Qu during the ], reported the discovery of dragon bones at Wucheng in ] Province.<ref>{{cite book|author=]|year=1992|title=Dinosaurian Faunas of China|publisher=China Ocean Press, Beijing|id=ISBN 3-540-52084-8}}</ref> In Europe, dinosaur fossils were generally believed to be the remains of ]s and other creatures killed by the ].


From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the ] in 1971. It included a '']'' attacking a '']'',<ref name=AMNH/> providing evidence that dinosaurs did indeed attack each other.<ref name=carpenter1998/> Additional evidence for attacking live prey is the partially healed tail of an '']'', a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten by a tyrannosaur but survived.<ref name=carpenter1998/> ] amongst some species of dinosaurs was confirmed by tooth marks found in ] in 2003, involving the theropod '']''.<ref name=rogersetal2003/>
]]]
'']'' was the first dinosaur to be formally described, in 1677, when part of a bone was recovered from a ] ] at ] near ], ]. This bone fragment was identified correctly as the lower extremity of the ] of an animal larger than anything living in modern times. The second dinosaur species to be identified, '']'', was discovered in 1822 by the English geologist ], who recognized similarities between his fossils and the bones of modern ]s. Two years later, the Rev ], a professor of ] at ], unearthed more fossilized bones of ''Megalosaurus'' and became the first person to describe dinosaurs in a ].


Comparisons between the ] of dinosaurs and modern birds and reptiles have been used to infer daily activity patterns of dinosaurs. Although it has been suggested that most dinosaurs were active during the day, these comparisons have shown that small predatory dinosaurs such as dromaeosaurids, '']'', and '']'' were likely ]. Large and medium-sized herbivorous and omnivorous dinosaurs such as ceratopsians, sauropodomorphs, hadrosaurids, ornithomimosaurs may have been ], active during short intervals throughout the day, although the small ornithischian '']'' was inferred to be ].<ref name=SchmitzMotani2011/>
The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English paleontologist ] coined the term "dinosaur". He recognized that the remains that had been found so far, ''Iguanodon'', ''Megalosaurus'' and '']'', shared a number of distinctive features, and so decided to present them as a distinct taxonomic group. With the backing of ], the husband of ], Owen established the ] in ], ], to display the national collection of dinosaur fossils and other biological and geological exhibits.


Based on fossil evidence from dinosaurs such as '']'', some ornithischian species seem to have led a partially ] (burrowing) lifestyle.<ref name=VMK07/> Many modern birds are ] (tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines.<ref>{{harvnb|Chiappe|Witmer|2002}}</ref> While some early bird-like species may have already been arboreal as well (including dromaeosaurids) such as ''Microraptor''<ref name=chatterjee2007/>) most non-avialan dinosaurs seem to have relied on land-based locomotion. A good understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior; the science of biomechanics, pioneered by ], has provided significant insight in this area. For example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast dinosaurs could run,<ref name=alexander2006/> whether ]s could create ]s via ]-like tail snapping,<ref name=goriely/> and whether sauropods could float.<ref name=Henderson2006/>
In 1858, the first known American dinosaur was discovered, in ] pits in the small town of ] (although fossils had been found before, their nature had not been correctly discerned). The creature was named ''] foulkii'', after the town and the discoverer, ]. It was an extremely important find; ''Hadrosaurus'' was the first nearly complete dinosaur skeleton found and it was clearly a ] creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of dinosaur mania in the ].


===Communication===
], (19th century photograph).]]
Modern birds ] by visual and auditory signals, and the wide diversity of visual display structures among fossil dinosaur groups, such as horns, frills, crests, sails, and feathers, suggests that visual communication has always been important in dinosaur biology.<ref name=senter2008/> Reconstruction of the plumage color of ''Anchiornis'' suggest the importance of color in visual communication in non-avian dinosaurs.<ref name="Li2010"/> Vocalization in non-avian dinosaurs is less certain. In birds, the ] plays no role in sound production. Instead, birds vocalize with a novel organ, the ], farther down the trachea.<ref name=":22"/> The earliest remains of a syrinx were found in a specimen of the duck-like '']'' dated 69 –66&nbsp;million years ago, and this organ is unlikely to have existed in non-avian dinosaurs.<ref name="Clarke2016"/>
], (19th century photograph).]]
]''. The crest may also have acted as a resonating chamber for sounds.]]


On the basis that non-avian dinosaurs did not have syrinxes and that their next close living relatives, crocodilians, use the larynx, Phil Senter, a paleontologist, has suggested that the non-avians could not vocalize, because the common ancestor would have been mute. He states that they mostly on visual displays and possibly non-vocal sounds, such as hissing, jaw-grinding or -clapping, splashing, and wing-beating (possible in winged maniraptoran dinosaurs).<ref name=senter2008/> Other researchers have countered that vocalizations also exist in turtles, the closest relatives of archosaurs, suggesting that the trait is ancestral to their lineage. In addition, vocal communication in dinosaurs is indicated by the development of advanced hearing in nearly all major groups. Hence the syrinx may have supplemented and then replaced the larynx as a vocal organ, without a "silent period" in bird evolution.<ref name=":0" />
Dinosaur mania was exemplified by the fierce rivalry between ] and ], both of whom raced to be the first to find new dinosaurs in what came to be known as the ]. The feud probably originated when Marsh publicly pointed out that Cope's reconstruction of an '']'' skeleton was flawed; Cope had inadvertently placed the ]'s head at what should have been the animal's tail end. The fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Marsh won the contest primarily because he was better funded through a relationship with the ]. Unfortunately, many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods; for example, their diggers often used ] to unearth bones (a method modern paleontologists would find appalling). Despite the pair's unrefined methods, their contributions to paleontology were vast; Marsh unearthed 86 new species of dinosaur and Cope discovered 56, for a total of 142 new species. Cope's collection is now at the ] in ], while Marsh's is on display at the ] at ].<ref>Williams, P. (1997). The Battle of the Bones. ''Dinosaur Cards''. Orbis Publishing Ltd. D36040607.</ref>


In 2023, a fossilized larynx was described, from a specimen of the ankylosaurid '']''. The structure was composed of ] and ]s, similar to those of non-avian reptiles; but the mobile cricoid–arytenoid joint and long arytenoid cartilages would have allowed air-flow control similar to that of birds, and thus could have made bird-like vocalizations. In addition, the cartilages were ], implying that laryngeal ossification is a feature of some non-avian dinosaurs.<ref name=":1" /> A 2016 study concludes that some dinosaurs may have produced closed-mouth vocalizations, such as cooing, hooting, and booming. These occur in both reptiles and birds and involve inflating the esophagus or tracheal pouches. Such vocalizations evolved independently in extant archosaurs numerous times, following increases in body size.<ref name="Tobias2016"/> The crests of some hadrosaurids and the nasal chambers of ankylosaurids may have been ].<ref name="Weishampel981"/><ref name="Witmer"/>
Since 1897, the search for dinosaur fossils has extended to every continent, including ]. The first Antarctic dinosaur to be discovered, the ]id '']'', was found on ] in 1986, although it was 1994 before an Antarctic species, the theropod '']'', was formally named and described in a scientific journal.


===Reproductive biology===
Current dinosaur "hot spots" include southern South America (especially ]) and ]. China in particular has produced many exceptional ] specimens due to the unique geology of its dinosaur beds, as well as an ancient arid climate particularly conducive to ]ization.
{{see also|Dinosaur egg}}
] ('']'')]]


All dinosaurs laid ]s. Dinosaur eggs were usually laid in a nest. Most species create somewhat elaborate nests which can be cups, domes, plates, beds scrapes, mounds, or burrows.<ref name="Hansell">{{harvnb|Hansell|2000}}</ref> Some species of modern bird have no nests; the cliff-nesting ] lays its eggs on bare rock, and male ]s keep eggs between their body and feet. Primitive birds and many non-avialan dinosaurs often lay eggs in communal nests, with males primarily incubating the eggs. While modern birds have only one functional ] and lay one egg at a time, more primitive birds and dinosaurs had two oviducts, like crocodiles. Some non-avialan dinosaurs, such as '']'', exhibited iterative laying, where the adult might lay a pair of eggs every one or two days, and then ensured simultaneous hatching by delaying ] until all eggs were laid.<ref name="Varrichioetal.02">{{cite journal |last1=Varricchio |first1=David J. |last2=Horner |first2=John R. |last3=Jackson |first3=Frankie D. |year=2002 |title=Embryos and eggs for the Cretaceous theropod dinosaur ''Troodon formosus'' |journal=Journal of Vertebrate Paleontology |location=Milton Park, Oxfordshire |publisher=Taylor & Francis for the Society of Vertebrate Paleontology |volume=22 |issue=3 |pages=564–576 |doi=10.1671/0272-4634(2002)0222.0.CO;2 |s2cid=85728452 |issn=0272-4634}}</ref>
==In popular culture==
].]]


When laying eggs, females grow a special type of bone between the hard outer bone and the ] of their limbs. This medullary bone, which is rich in ], is used to make eggshells. A discovery of features in a ''Tyrannosaurus'' skeleton provided evidence of medullary bone in extinct dinosaurs and, for the first time, allowed paleontologists to establish the sex of a fossil dinosaur specimen. Further research has found medullary bone in the carnosaur ''Allosaurus'' and the ornithopod '']''. Because the line of dinosaurs that includes ''Allosaurus'' and ''Tyrannosaurus'' diverged from the line that led to ''Tenontosaurus'' very early in the evolution of dinosaurs, this suggests that the production of medullary tissue is a general characteristic of all dinosaurs.<ref name=LW08/>
{{main|Dinosaurs in popular culture}}
] '']'' at the ]. ]]
By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured people's imagination and become an enduring part of human popular culture. Dinosaur exhibitions, parks and museum exhibits around the world both cater to and reinforce the public's interest. The popular preoccupation with dinosaurs is also reflected in a broad array of ] and ] works.
Another widespread trait among modern birds (but see below in regards to fossil groups and extant ]) is parental care for young after hatching. ] 1978 discovery of a '']'' ("good mother lizard") nesting ground in Montana demonstrated that parental care continued long after birth among ornithopods.<ref name=HM79/> A specimen of the ] '']'' was discovered in a ] in 1993,<ref name="search.eb"/> which may indicate that they had begun using an insulating layer of feathers to keep the eggs warm.<ref>{{harvnb|Currie|Koppelhus|Shugar|Wright|2004|pp=|loc=chpt. 11: "Dinosaur Brooding Behavior and the Origin of Flight Feathers" by Thomas P. Hopp and Mark J. Orsen.}}</ref> An embryo of the basal sauropodomorph '']'' was found without teeth, indicating that some parental care was required to feed the young dinosaurs.<ref name=Reiszetal05/> Trackways have also confirmed parental behavior among ornithopods from the ] in northwestern ].<ref name=BBCtracks/>


However, there is ample evidence of ] or ] among many dinosaur species, particularly theropods. For instance, non-] birds have been abundantly demonstrated to have had slow growth rates, ]-like egg burying behavior and the ability to fly soon after birth.<ref>{{cite journal |first1=Zhonghe |last1=Zhou |first2=Fucheng |last2=Zhang |year=2004 |title=A Precocial Avian Embryo from the Lower Cretaceous of China |journal=Science |location=Washington, D.C. |publisher=American Association for the Advancement of Science |volume=306 |issue=5696 |page=653 |doi=10.1126/science.1100000 |issn=0036-8075 |pmid=15499011|s2cid=34504916 }}</ref><ref>{{cite journal |url=https://blogs.scientificamerican.com/tetrapod-zoology/drowned-cretaceous-bird-colony/ |url-status=live |title=A drowned nesting colony of Late Cretaceous birds |last=Naish |first=Darren |author-link=Darren Naish |date=May 15, 2012 |journal=Science |volume=306 |issue=5696 |page=653 |publisher=] |doi=10.1126/science.1100000 |pmid=15499011 |s2cid=34504916 |archive-url=https://web.archive.org/web/20180925031243/https://blogs.scientificamerican.com/tetrapod-zoology/drowned-cretaceous-bird-colony/ |archive-date=September 25, 2018 |access-date=November 16, 2019}}</ref><ref>{{cite journal |last1=Fernández |first1=Mariela S. |last2=García |first2=Rodolfo A. |last3=Fiorelli |first3=Lucas |last4=Scolaro |first4=Alejandro |last5=Salvador |first5=Rodrigo B. |last6=Cotaro |first6=Carlos N. |last7=Kaiser |first7=Gary W. |last8=Dyke |first8=Gareth J. |author8-link=Gareth J. Dyke |display-authors=3 |year=2013 |title=A Large Accumulation of Avian Eggs from the Late Cretaceous of Patagonia (Argentina) Reveals a Novel Nesting Strategy in Mesozoic Birds |journal=PLOS ONE |location=San Francisco, CA |publisher=PLOS |volume=8 |issue=4 |page=e61030 |doi=10.1371/journal.pone.0061030 |pmid=23613776 |pmc=3629076 |bibcode=2013PLoSO...861030F |issn=1932-6203|doi-access=free }}</ref><ref>{{cite journal |last1=Deeming |first1=Denis Charles |last2=Mayr |first2=Gerald |author2-link=Gerald Mayr |date=May 2018 |title=Pelvis morphology suggests that early Mesozoic birds were too heavy to contact incubate their eggs |journal=] |location=Hoboken, NJ |publisher=Wiley-Blackwell on behalf of the ] |volume=31 |issue=5 |pages=701–709 |doi=10.1111/jeb.13256 |pmid=29485191 |s2cid=3588317 |issn=1010-061X|url=http://eprints.lincoln.ac.uk/id/eprint/31436/13/31436%2031291%20Deeming_et_al-2018-Journal_of_Evolutionary_Biology.pdf |archive-url=https://web.archive.org/web/20200602094641/http://eprints.lincoln.ac.uk/id/eprint/31436/13/31436%2031291%20Deeming_et_al-2018-Journal_of_Evolutionary_Biology.pdf |archive-date=2020-06-02 |url-status=live }}</ref> Both ''Tyrannosaurus'' and ''Troodon'' had juveniles with clear superprecociality and likely occupying different ecological niches than the adults.<ref name="Varrichioetal.02"/> Superprecociality has been inferred for sauropods.<ref>{{cite journal |last1=Myers |first1=Timothy S. |last2=Fiorillo |first2=Anthony R. |year=2009 |title=Evidence for gregarious behavior and age segregation in sauropod dinosaurs |journal=] |location=Amsterdam |publisher=Elsevier |volume=274 |issue=1–2 |pages=96–104 |doi=10.1016/j.palaeo.2009.01.002 |issn=0031-0182|bibcode=2009PPP...274...96M |url=http://doc.rero.ch/record/16633/files/PAL_E962.pdf |archive-url=https://web.archive.org/web/20200529011847/http://doc.rero.ch/record/16633/files/PAL_E962.pdf |archive-date=2020-05-29 |url-status=live }}</ref>
Notable examples of older fictional works featuring dinosaurs include ]'s book '']''; the 1933 film '']''; and '']''.


Genital structures are unlikely to fossilize as they lack scales that may allow preservation via pigmentation or residual calcium phosphate salts. In 2021, the best preserved specimen of a dinosaur's ] exterior was described for ''Psittacosaurus'', demonstrating lateral swellings similar to crocodylian musk glands used in social displays by both sexes and pigmented regions which could also reflect a signalling function. However, this specimen on its own does not offer enough information to determine whether this dinosaur had sexual signalling functions; it only supports the possibility. Cloacal visual signalling can occur in either males or females in living birds, making it unlikely to be useful to determine sex for extinct dinosaurs.<ref name="cloacal-vent">{{cite journal|doi=10.1016/j.cub.2020.12.039|title=A cloacal opening in a non-avian dinosaur|first1=Jakob|last1=Vinther|first2=Robert|last2=Nicholls|first3=Diane A.|last3=Kelly|journal=Current Biology|volume=31|pages=R1–R3|date=February 22, 2021|issue=4|publisher=Elsevier|pmid=33472049|s2cid=231644183|doi-access=free|bibcode=2021CBio...31.R182V }}</ref>
==Religious views==
{{main|Religious perspectives on dinosaurs}}
Various religious groups have views about dinosaurs that differ from those held by scientists. While many mainstream scientists respect these views as ] positions, they argue that religiously-inspired interpretations of dinosaurs do not withstand serious ]. See the referenced article for specific examples and further context.


==See also== ===Physiology===
{{Main|Physiology of dinosaurs}}
*]
Because both modern crocodilians and birds have four-chambered hearts (albeit modified in crocodilians), it is likely that this is a trait shared by all archosaurs, including all dinosaurs.<ref name=CH04/> While all modern birds have high metabolisms and are ]ic ("warm-blooded"), a vigorous debate has been ongoing since the 1960s regarding how far back in the dinosaur lineage this trait extended. Various researchers have supported dinosaurs as being endothermic, ]ic ("cold-blooded"), or somewhere in between.<ref name="pontzer2009">{{cite journal |last1=Pontzer |first1=H. |last2=Allen |first2=V. |last3=Hutchinson |first3=J.R. |year=2009 |title=Biomechanics of running indicates endothermy in bipedal dinosaurs |journal=PLOS ONE |volume=4 |issue=11 |page=e7783 |doi=10.1371/journal.pone.0007783 |doi-access=free |pmc=2772121 |pmid=19911059 |bibcode=2009PLoSO...4.7783P |issn=1932-6203}}</ref> An emerging consensus among researchers is that, while different lineages of dinosaurs would have had different metabolisms, most of them had higher metabolic rates than other reptiles but lower than living birds and mammals,<ref name="benson2018">{{cite journal |last1=Benson |first1=R.B.J. |year=2018 |title=Dinosaur Macroevolution and Macroecology |journal=Annual Review of Ecology, Evolution, and Systematics |volume=49 |pages=379–408 |doi=10.1146/annurev-ecolsys-110617-062231|s2cid=92837486 |doi-access=free }}</ref> which is termed ]y by some.<ref name="grady2014">{{cite journal |last1=Grady |first1=J.M. |last2=Enquist |first2=B.J. |last3=Dettweiler-Robinson |first3=E. |last4=Wright |first4=N.A. |last5=Smith |first5=F.A. |year=2014 |title=Evidence for mesothermy in dinosaurs |journal=Science |volume=344 |issue=6189 |pages=1268–1272 |doi=10.1126/science.1253143|pmid=24926017 |bibcode=2014Sci...344.1268G |s2cid=9806780 }}</ref> Evidence from crocodiles and their extinct relatives suggests that such elevated metabolisms could have developed in the earliest archosaurs, which were the common ancestors of dinosaurs and crocodiles.<ref name="legendre2016">{{cite journal |last1=Legendre |first1=L.J. |last2=Guénard |first2=G. |last3=Botha-Brink |first3=J. |last4=Cubo |first4=J. |title=Palaeohistological Evidence for Ancestral High Metabolic Rate in Archosaurs |journal=Systematic Biology |year=2016 |volume=65 |issue=6 |pages=989–996 |doi=10.1093/sysbio/syw033|pmid=27073251 |doi-access=free }}</ref><ref name="seymour2004">{{cite journal |last1=Seymour |first1=R.S. |last2=Bennett-Stamper |first2=C.L. |last3=Johnston |first3=S.D. |last4=Carrier |first4=D.R. |last5=Grigg |first5=G.C. |year=2004 |title=Evidence for endothermic ancestors of crocodiles at the stem of archosaur evolution |journal=Physiological and Biochemical Zoology |volume=77 |issue=6 |pages=1051–1067 |doi=10.1093/sysbio/syw033|pmid=27073251 |doi-access=free }}</ref>
*]s
*]
*]
*]s
* ] - The Senckenberg Museum in ], ] boasts the largest exhibition of large dinosaurs in ].


]'' as an aquatic, tail-dragging animal, by ], typified early views on dinosaur lifestyles.]]
==Notes and references==
After non-avian dinosaurs were discovered, paleontologists first posited that they were ectothermic. This was used to imply that the ancient dinosaurs were relatively slow, sluggish organisms, even though many modern reptiles are fast and light-footed despite relying on external sources of heat to regulate their body temperature. The idea of dinosaurs as ectothermic remained a prevalent view until ], an early proponent of dinosaur endothermy, published an influential paper on the topic in 1968. Bakker specifically used anatomical and ecological evidence to argue that sauropods, which had hitherto been depicted as sprawling aquatic animals with their tails dragging on the ground, were endotherms that lived vigorous, terrestrial lives. In 1972, Bakker expanded on his arguments based on energy requirements and predator-prey ratios. This was one of the seminal results that led to the dinosaur renaissance.<ref name="bakker1968">{{cite journal |last1=Bakker |first1=R.T. |author-link=Robert T. Bakker |year=1968 |title=The Superiority of Dinosaurs |journal=Discovery: Magazine of the Peabody Museum of Natural History |volume=3 |issue=2 |pages=11–22 |issn=0012-3625 |oclc=297237777}}</ref><ref name="bakker1972">{{cite journal |last1=Bakker |first1=R.T. |author-link=Robert T. Bakker |year=1972 |title=Anatomical and Ecological Evidence of Endothermy in Dinosaurs |journal=Nature |volume=238 |issue=5359 |pages=81–85 |doi=10.1038/238081a0|bibcode=1972Natur.238...81B |s2cid=4176132 }}</ref><ref name="taylor2010"/><ref name="parsons2001"/>
<div class="references-small">
<!-- Dead note "reptilia": From the classical standpoint, reptiles included all the amniotes except birds and mammals. Thus reptiles were defined as the set of animals that includes crocodiles, alligators, tuatara, lizards, snakes, amphisbaenians and turtles, grouped together as the class Reptilia. However, many taxonomists have begun to insist that taxa should be monophyletic, that is, groups should include all descendants of a particular form. The reptiles as defined here would be paraphyletic, since they exclude both birds and mammals, although these also developed from the original reptile. Thus, some cladists redefine Reptilia as a monophyletic group, including both the classic reptiles as well as the birds and perhaps the mammals (depending on ideas about their relationships). Others abandon it as a formal taxon altogether, dividing it into several different classes. -->
<references />
<!-- Dead note "jpii": Catholic Opinions on Evolutionary Origins. -->


One of the greatest contributions to the modern understanding of dinosaur physiology has been ], the study of microscopic tissue structure in dinosaurs.<ref name="erickson2014">{{cite journal |last1=Erickson |first1=G.M. |year=2014 |title=On dinosaur growth |journal=Annual Review of Earth and Planetary Sciences |volume=42 |issue=1 |pages=675–697 |doi=10.1146/annurev-earth-060313-054858|bibcode=2014AREPS..42..675E }}</ref><ref name="bailleul2019">{{cite journal |last1=Bailleul |first1=A.M. |last2=O'Connor |first2=J. |last3=Schweitzer |first3=M.H. |year=2019 |title=Dinosaur paleohistology: review, trends and new avenues of investigation |journal=PeerJ |volume=7 |page=e7764 |doi=10.7717/peerj.7764|pmid=31579624 |pmc=6768056 |doi-access=free }}</ref> From the 1960s forward, ] suggested that the presence of fibrolamellar bone—bony tissue with an irregular, fibrous texture and filled with blood vessels—was indicative of consistently fast growth and therefore endothermy. Fibrolamellar bone was common in both dinosaurs and pterosaurs,<ref name="dericqles1974">{{cite journal |last1=De Ricqlès |first1=A. |year=1974 |title=Evolution of endothermy: histological evidence |journal=Evolutionary Theory |volume=1 |issue=2 |pages=51–80 |url=http://www.stuartsumida.com/BIOL622/Ricqles1974.pdf |archive-url=https://web.archive.org/web/20210417055907/http://www.stuartsumida.com/BIOL622/Ricqles1974.pdf |archive-date=2021-04-17 |url-status=live}}</ref><ref name="dericqles1980">{{cite book |last1=De Ricqlès |first1=A. |year=1980 |chapter=Tissue structures of dinosaur bone, functional significance and possible relation to dinosaur physiology |editor-last1=Thomas |editor-first1=R.D.K. |editor-last2=Olson |editor-first2=E.C. |title=A Cold Look at the Warm-Blooded Dinosaurs |location=New York |publisher=American Association for the Advancement of Science |pages=103–139}}</ref> though not universally present.<ref name="padian2004">{{cite journal |last1=Padian |first1=K. |last2=Horner |first2=J.R. |last3=de Ricqlès |first3=A. |year=2004 |title=Growth in small dinosaurs and pterosaurs: the evolution of archosaurian growth strategies |journal=Journal of Vertebrate Paleontology |volume=24 |issue=3 |pages=555–571 |doi=10.1671/0272-4634(2004)0242.0.CO;2|s2cid=86019906 |url=http://doc.rero.ch/record/15191/files/PAL_E2467.pdf }}</ref><ref name="desouza2020">{{cite journal |last1=de Souza |first1=G.A. |last2=Bento Soares |first2=M. |last3=Souza Brum |first3=A. |last4=Zucolotto |first4=M. |last5=Sayão |first5=J.M. |last6=Carlos Weinschütz |first6=L. |last7=Kellner |first7=A.W.A. |year=2020 |title=Osteohistology and growth dynamics of the Brazilian noasaurid ''Vespersaurus paranaensis'' Langer et al., 2019 (Theropoda: Abelisauroidea) |journal=PeerJ |volume=8 |pages=e9771 |doi=10.7717/peerj.9771 |pmid=32983636 |pmc=7500327 |s2cid=221906765 |doi-access=free }}</ref> This has led to a significant body of work in reconstructing ] and modeling the evolution of growth rates across various dinosaur lineages,<ref name="physiologyrefs">For examples of this work conducted on different dinosaur lineages, see
</div>


* {{cite journal |last1=Erickson |first1=G.M. |last2=Tumanova |first2=T.A. |year=2000 |title=Growth curve of ''Psittacosaurus mongoliensis'' Osborn (Ceratopsia: Psittacosauridae) inferred from long bone histology |journal=Zoological Journal of the Linnean Society |volume=130 |issue=4 |pages=551–566 |doi=10.1111/j.1096-3642.2000.tb02201.x |s2cid=84241148}}
==General references==
* {{cite journal |last1=Erickson |first1=G. |last2=Rogers |first2=K. |last3=Yerby |first3=S. |year=2001 |title=Dinosaurian growth patterns and rapid avian growth rates |journal=Nature |volume=412 |issue=429–433 |pages=429–433 |bibcode=2001Natur.412..429E |doi=10.1038/35086558 |pmid=11473315 |s2cid=4319534}}{{Erratum|doi=10.1038/nature16488|pmid=26675731|http://retractionwatch.com/2016/03/01/high-profile-critic-slams-nature-letters-about-dinosaur-growth-following-corrections/ ''Retraction Watch''|checked=yes}}
<div class="references-small">
* {{cite journal |last1=Erickson |first1=G. |last2=Makovicky |first2=P. |last3=Currie |first3=P. |last4=Norell |first4=M.A. |last5=Yerby |first5=S.A. |last6=Brochu |first6=C.A. |year=2004 |title=Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs |url=http://doc.rero.ch/record/15279/files/PAL_E2578.pdf |url-status=live |journal=Nature |volume=430 |issue=7001 |pages=772–775 |bibcode=2004Natur.430..772E |doi=10.1038/nature02699 |pmid=15306807 |s2cid=4404887 |archive-url=https://web.archive.org/web/20200714024211/http://doc.rero.ch/record/15279/files/PAL_E2578.pdf |archive-date=2020-07-14}}{{Erratum|doi=10.1038/nature16487|pmid=26675726|http://retractionwatch.com/2016/03/01/high-profile-critic-slams-nature-letters-about-dinosaur-growth-following-corrections/ ''Retraction Watch''|checked=yes}}
* Kevin Padian, and Philip J. Currie. (1997). ''Encyclopedia of Dinosaurs''. Academic Press. ISBN 0-12-226810-5. (Articles are written by experts in the field).
* {{cite journal |last1=Lehman |first1=T.M. |last2=Woodward |first2=H.N. |year=2008 |title=Modeling growth rates for sauropod dinosaurs |url=http://doc.rero.ch/record/16723/files/PAL_E3766.pdf |journal=Paleobiology |volume=34 |issue=2 |pages=264–281 |doi=10.1666/0094-8373(2008)0342.0.CO;2 |s2cid=84163725}}
* ] (2000). ''The Scientific American Book of Dinosaurs''. St. Martin's Press. ISBN 0-312-26226-4.
* {{cite journal |last1=Horner |first1=J.R. |last2=de Ricqles |first2=A. |last3=Padian |first3=K. |last4=Scheetz |first4=R.D. |year=2009 |title=Comparative long bone histology and growth of the "hypsilophodontid" dinosaurs ''Orodromeus makelai'', ''Dryosaurus altus'', and ''Tenontosaurus tillettii'' (Ornithischia: Euornithopoda) |journal=Journal of Vertebrate Paleontology |volume=29 |issue=3 |pages=734–747 |bibcode=2009JVPal..29..734H |doi=10.1671/039.029.0312 |s2cid=86277619}}
*Paul, Gregory S. (2002). ''Dinosaurs of the Air: The Evolution and Loss of flight in Dinosaurs and Birds''. Baltimore: The Johns Hopkins University Press. ISBN 0-8018-6763-0.
* {{cite journal |last1=Woodward |first1=H. |last2=Freedman Fowler |first2=E. |last3=Farlow |first3=J. |last4=Horner |first4=J. |year=2015 |title=''Maiasaura'', a model organism for extinct vertebrate population biology: A large sample statistical assessment of growth dynamics and survivorship |journal=Paleobiology |volume=41 |issue=4 |pages=503–527 |bibcode=2015Pbio...41..503W |doi=10.1017/pab.2015.19 |s2cid=85902880}}</ref> which has suggested overall that dinosaurs grew faster than living reptiles.<ref name="bailleul2019"/> Other lines of evidence suggesting endothermy include the presence of feathers and other types of body coverings in many lineages (see {{section link||Feathers}}); more consistent ratios of the isotope ] in bony tissue compared to ectotherms, particularly as latitude and thus air temperature varied, which suggests stable internal temperatures<ref name="amiot2006">{{cite journal |last1=Amiot |first1=R. |last2=Lécuyer |first2=C. |last3=Buffetaut |first3=E. |last4=Escarguel |first4=G. |last5=Fluteau |first5=F. |last6=Martineau |first6=F. |year=2006 |title=Oxygen isotopes from biogenic apatites suggest widespread endothermy in Cretaceous dinosaurs |journal=Earth and Planetary Science Letters |volume=246 |issue=1–2 |pages=41–54 |doi=10.1016/j.epsl.2006.04.018|bibcode=2006E&PSL.246...41A |url=http://doc.rero.ch/record/14262/files/PAL_E2159.pdf }}</ref><ref name="amiot2010">{{cite journal |last1=Amiot |first1=R. |last2=Wang |first2=X. |last3=Lécuyer |first3=C. |last4=Buffetaut |first4=E. |last5=Boudad |first5=L. |last6=Cavin |first6=L. |last7=Ding |first7=Z. |last8=Fluteau |first8=F. |last9=Kellner |first9=A.W.A. |last10=Tong |first10=H. |last11=Zhang |first11=F. |year=2010 |title=Oxygen and carbon isotope compositions of middle Cretaceous vertebrates from North Africa and Brazil: ecological and environmental significance |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=297 |issue=2 |pages=439–451 |doi=10.1016/j.palaeo.2010.08.027|bibcode=2010PPP...297..439A }}</ref> (although these ratios can be altered during fossilization<ref name="kolodny1996">{{cite journal |last1=Kolodny |first1=Y. |last2=Luz |first2=B. |last3=Sander |first3=M. |last4=Clemens |first4=W.A. |year=1996 |title=Dinosaur bones: fossils or pseudomorphs? The pitfalls of physiology reconstruction from apatitic fossils |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=126 |issue=1–2 |pages=161–171 |doi=10.1016/S0031-0182(96)00112-5|bibcode=1996PPP...126..161K |url=http://doc.rero.ch/record/14443/files/PAL_E1627.pdf }}</ref>); and the discovery of ], which lived in Australia, Antarctica, and Alaska when these places would have had cool, temperate climates.<ref name="paul1988">{{cite journal |last1=Paul |first1=G.S. |year=1988 |title=Physiological, migratorial, climatological, geophysical, survival, and evolutionary implications of Cretaceous polar dinosaurs |journal=Journal of Paleontology |volume=62 |issue=4 |pages=640–652 |jstor=1305468|ref=none}}</ref><ref name="clemens1993">{{cite journal |last1=Clemens |first1=W.A. |last2=Nelms |first2=L.G. |year=1993 |title=Paleoecological implications of Alaskan terrestrial vertebrate fauna in latest Cretaceous time at high paleolatitudes |journal=Geology |volume=21 |issue=6 |pages=503–506 |doi=10.1130/0091-7613(1993)021<0503:PIOATV>2.3.CO;2|bibcode=1993Geo....21..503C }}</ref><ref name="rich2002">{{cite journal |last1=Rich |first1=T.H. |last2=Vickers-Rich |first2=P. |last3=Gangloff |first3=R.A. |year=2002 |title=Polar dinosaurs |journal=Science |volume=295 |issue=5557 |pages=979–980 |doi=10.1126/science.1068920|pmid=11834803 |s2cid=28065814 }}</ref><ref name="buffetaut2004">{{cite journal |last1=Buffetaut |first1=E. |year=2004 |title=Polar dinosaurs and the question of dinosaur extinction: a brief review |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=214 |issue=3 |pages=225–231 |doi=10.1016/j.palaeo.2004.02.050|url=http://doc.rero.ch/record/15383/files/PAL_E2734.pdf |archive-url=https://web.archive.org/web/20200608120307/http://doc.rero.ch/record/15383/files/PAL_E2734.pdf |archive-date=2020-06-08 |url-status=live }}</ref>
*] (2004). ''The Dinosauria''. University of California Press; 2nd edition. ISBN 0-520-24209-2.
</div>


]s of an ] and a bird]]
==External links==
In saurischian dinosaurs, higher metabolisms were supported by the evolution of the avian respiratory system, characterized by an extensive system of ]s that extended the lungs and invaded many of the bones in the skeleton, making them hollow.<ref name=Sereno2008/> Such respiratory systems, which may have appeared in the earliest saurischians,<ref name="oconnor2009">{{cite journal |last1=O'Connor |first1=P.M. |year=2009 |title=Evolution of archosaurian body plans: skeletal adaptations of an air-sac-based breathing apparatus in birds and other archosaurs |journal=Journal of Experimental Zoology Part A: Ecological Genetics and Physiology |volume=311 |issue=8 |pages=629–646 |doi=10.1002/jez.548|pmid=19492308 |bibcode=2009JEZA..311..629O }}</ref> would have provided them with more oxygen compared to a mammal of similar size, while also having a larger resting ] and requiring a lower breathing frequency, which would have allowed them to sustain higher activity levels.<ref name="sander2011"/> The rapid airflow would also have been an effective cooling mechanism, which in conjunction with a lower metabolic rate<ref name="eagle2011">{{cite journal |last1=Eagle |first1=R.A. |last2=Tütken |first2=T. |last3=Martin |first3=T.S. |last4=Tripati |first4=A.K. |last5=Fricke |first5=H.C. |last6=Connely |first6=M. |last7=Cifelli |first7=R.L. |last8=Eiler |first8=J.M. |year=2011 |title=Dinosaur body temperatures determined from isotopic (<sup>13</sup>C-<sup>18</sup>O) ordering in fossil biominerals |journal=Science |volume=333 |issue=6041 |pages=443–445 |pmid=21700837 |doi=10.1126/science.1206196 |bibcode=2011Sci...333..443E |s2cid=206534244}}</ref> would have prevented large sauropods from overheating. These traits may have enabled sauropods to grow quickly to gigantic sizes.<ref name="wedel2003">{{cite journal |last1=Wedel |first1=M.J. |year=2003 |title=Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs |journal=Paleobiology |volume=29 |issue=2 |pages=243–255 |doi=10.1017/S0094837300018091|bibcode=2003Pbio...29..243W |url=http://doc.rero.ch/record/14872/files/PAL_E2010.pdf }}</ref><ref name="perry2009">{{cite journal |last1=Perry |first1=S.F. |last2=Christian |first2=A. |last3=Breuer |first3=T. |last4=Pajor |first4=N. |last5=Codd |first5=J.R. |year=2009 |title=Implications of an avian-style respiratory system for gigantism in sauropod dinosaurs |journal=Journal of Experimental Zoology Part A: Ecological Genetics and Physiology |volume=311 |issue=8 |pages=600–610 |doi=10.1002/jez.517 |pmid=19189317|bibcode=2009JEZA..311..600P }}</ref> Sauropods may also have benefitted from their size—their small surface area to volume ratio meant that they would have been able to thermoregulate more easily, a phenomenon termed ].<ref name="sander2011"/><ref name="alexander1998">{{cite journal |last1=Alexander |first1=R.M. |year=1998 |title=All-time giants: the largest animals and their problems |journal=Palaeontology |volume=41 |pages=1231–1245 |url=https://www.palass.org/publications/palaeontology-journal/archive/41/6/article_pp1231-1245}}</ref>
<!--Sorted (roughly) from least to most technical-->
{{commons|Dinosauria}}
{{Spoken Misplaced Pages|Dinosaur.ogg|2005-12-30}}
;For children
* From MantyWeb Educational Software. Kid's site, facts, games.
* From Yahooligans! Science. Glossaries, dino cards and indexes.
* From Enchanted Learning. Kid's site, info pages, theories, history.
* so the dinos get into your fingers, also for adults.
* Award-winning online children's science magazine.


Like other reptiles, dinosaurs are primarily ], that is, their ]s extract nitrogenous wastes from their bloodstream and excrete it as ] instead of ] or ] via the ureters into the intestine. This would have helped them to conserve water.<ref name="benson2018"/> In most living species, uric acid is excreted along with feces as a semisolid waste.<ref name="tsahar2005">{{cite journal |last1=Tsahar |first1=E. |last2=Martínez del Rio |first2=C. |last3=Izhaki |first3=I. |last4=Arad |first4=Z. |year=2005 |title=Can birds be ammonotelic? Nitrogen balance and excretion in two frugivores |journal=] |volume=208 |issue=6 |pages=1025–1034 |url= https://jeb.biologists.org/content/jexbio/208/6/1025.full.pdf |url-status=live |doi=10.1242/jeb.01495 |issn=0022-0949 |pmid=15767304 |s2cid=18540594 |archive-url=https://web.archive.org/web/20191017110333/https://jeb.biologists.org/content/jexbio/208/6/1025.full.pdf |archive-date=October 17, 2019 |access-date=October 31, 2019}}</ref><ref name="skadhauge2003">{{cite journal |last1=Skadhauge |first1=E. |last2=Erlwanger |first2=K.H. |last3=Ruziwa |first3=S.D. |last4=Dantzer |first4=V. |last5=Elbrønd |first5=V.S. |last6=Chamunorwa |first6=J.P. |year=2003 |title=Does the ostrich (''Struthio camelus'') coprodeum have the electrophysiological properties and microstructure of other birds? |journal=] |volume=134 |issue=4 |pages=749–755 |doi=10.1016/S1095-6433(03)00006-0 |issn=1095-6433 |pmid=12814783}}</ref> However, at least some modern birds (such as ]s) can be facultatively ], excreting most of the nitrogenous wastes as ammonia.<ref name="preest1997">{{cite journal |last1=Preest |first1=M.R. |last2=Beuchat |first2=C.A. |year=1997 |title=Ammonia excretion by hummingbirds |journal=Nature |volume=386 |issue=6625 |pages=561–562 |bibcode=1997Natur.386..561P |doi=10.1038/386561a0 |s2cid=4372695 |issn=0028-0836}}</ref> This material, as well as the output of the intestines, emerges from the ].<ref name="mora1965">{{cite journal |last1=Mora |first1=J. |last2=Martuscelli |first2=J. |last3=Ortiz Pineda |first3=J. |last4=Soberon |first4=G. |year=1965 |title=The Regulation of Urea-Biosynthesis Enzymes in Vertebrates |journal=] |volume=96 |issue=1 |pages=28–35 |issn=0264-6021 |doi=10.1042/bj0960028 |pmc=1206904 |pmid=14343146}}</ref><ref name="packard1966">{{cite journal |last=Packard |first=G.C. |year=1966 |title=The Influence of Ambient Temperature and Aridity on Modes of Reproduction and Excretion of Amniote Vertebrates |journal=The American Naturalist |volume=100 |issue=916 |pages=667–682 |doi=10.1086/282459 |issn=0003-0147 |jstor=2459303 |s2cid=85424175}}</ref> In addition, many species regurgitate ],<ref name="balgooyen1971">{{cite journal |last=Balgooyen |first=T.G. |year=1971 |title=Pellet Regurgitation by Captive Sparrow Hawks (''Falco sparverius'') |journal=] |volume=73 |issue=3 |pages=382–385 |jstor=1365774 |url=https://sora.unm.edu/sites/default/files/journals/condor/v073n03/p0382-p0385.pdf |doi=10.2307/1365774 |archive-url=https://web.archive.org/web/20190404001957/https://sora.unm.edu/sites/default/files/journals/condor/v073n03/p0382-p0385.pdf |archive-date=April 4, 2019 |access-date=October 30, 2019}}</ref> and fossil pellets are known as early as the Jurassic from '']''.<ref name="xu2018">{{cite journal |last1=Xu |first1=X. |last2=Li |first2=F. |last3=Wang |first3=Y. |last4=Sullivan |first4=C. |last5=Zhang |first5=F. |last6=Zhang |first6=X. |last7=Sullivan |first7=C. |last8=Wang |first8=X. |last9=Zheng |first9=X. |year=2018 |title=Exceptional dinosaur fossils reveal early origin of avian-style digestion |journal=Scientific Reports |volume=8 |issue=1 |pages=14217 |doi=10.1038/s41598-018-32202-x |pmid=30242170 |issn=2045-2322 |pmc=6155034 |bibcode=2018NatSR...814217Z}}</ref>
;Popular
* From the ]. London popular site, well illustrated dino directory.
* Humorous educational video about the dinosaur basics.
* The Dino-headlines from around the world. Recent news on dinosaurs, including finds and discoveries, lots of links.
* The Discoveries of Dr. Joan Wiffen, New Zealand's Dinosaur Lady
* Timeline of the discovery of Dinosaurs.
* From the ]. Popular overview.
* From the ]. Popular site, very well illustrated.
* From DinoData. Summaries of modern debates about dinosaurs.
* From UC Berkeley Museum of Paleontology Detailed information - scroll down for menu.
* A gallery of dino-paintings.
*
* All about Dinosaurs with current featured articles.


{{anchor|intelligence}}The size and shape of the brain can be partly reconstructed based on the surrounding bones. In 1896, Marsh calculated ratios between brain weight and body weight of seven species of dinosaurs, showing that the brain of dinosaurs was proportionally smaller than in today's crocodiles, and that the brain of ''Stegosaurus'' was smaller than in any living land vertebrate. This contributed to the widespread public notion of dinosaurs as being sluggish and extraordinarily stupid. Harry Jerison, in 1973, showed that proportionally smaller brains are expected at larger body sizes, and that brain size in dinosaurs was not smaller than expected when compared to living reptiles.<ref name="russell1997"/> Later research showed that relative brain size progressively increased during the evolution of theropods, with the highest intelligence – comparable to that of modern birds – calculated for the troodontid ''Troodon''.<ref name="brusatte2012"/>
;Technical
*
* From PaleoClones. Current dino news.
* From '']''. Article on the rapid extinction of dinosaurs.
* From the '']''. Articles, latest news but out of date.
* From Coquina Press. Online technical journal.
* Article from by Z.K. Silagadze.


==Origin of birds==
;Very technical
{{Main|Origin of birds}}
* A searchable dinosaur database, from the University of Bristol, with dinosaur lists, classification, pictures, and more.
* Technical site, essays, classification, anatomy.
* Technical site, essays, pronunciation, dictionary.
* By T. Michael Keesey. Technical site, cladogram, illustrations and animations.
* By Justin Tweet. Includes a cladogram and small essays on each relevant genera and species.
* From . A detailed and wonderful amateur site about all things paleo.
* A very extensive site regarding dinosaur information.


The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by ].<ref name=huxley1868/> After the work of ] in the early 20th century, the ] of birds as dinosaur descendants was abandoned in favor of the idea of them being descendants of generalized ], with the key piece of evidence being the supposed lack of ]s in dinosaurs.<ref name=heilmann/> However, as later discoveries showed, clavicles (or a single fused ], which derived from separate clavicles) were not actually absent;<ref name=KP04/> they had been found as early as 1924 in '']'', but misidentified as an ].<ref name=HO24/> In the 1970s, Ostrom revived the dinosaur–bird theory,<ref name=ostrom1973/> which gained momentum in the coming decades with the advent of cladistic analysis,<ref name=gauthier1986/> and a great increase in the discovery of small theropods and early birds.<ref name=TRHJ00/> Of particular note have been the fossils of the Jehol Biota, where a variety of theropods and early birds have been found, often with feathers of some type.<ref name="NYT-20161208" /><ref name=KP04/> Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are now generally accepted to have been their closest ancient relatives.<ref name=Mayretal2005/> They are most closely allied with maniraptoran coelurosaurs.<ref name=KP04/> A minority of scientists, most notably ] and ], have proposed other evolutionary paths, including revised versions of Heilmann's basal archosaur proposal,<ref name=martin2004/> or that maniraptoran theropods are the ancestors of birds but themselves are not dinosaurs, only ] with dinosaurs.<ref name=AF02/>
;Bird-dinosaur and dinosaur warm-bloodedness discussion
* Are birds Dinosaurs?
* Site focusing on the dinosaur-bird relationship.
*
*
*
*


===Feathers===
{{featured article}}
{{Main|Feathered dinosaurs}}
]'', '']'', '']'' and '']'']]
Feathers are one of the most recognizable characteristics of modern birds, and a trait that was also shared by several non-avian dinosaurs. Based on the current distribution of fossil evidence, it appears that feathers were an ancestral dinosaurian trait, though one that may have been selectively lost in some species.<ref name=switeknature>{{cite journal |url=https://www.nature.com/articles/nature.2012.10933 |title=Rise of the fuzzy dinosaurs |last1=Switek |first1=Brian |date=July 2, 2012 |department=News |journal=Nature |publisher=Nature Research |location=London |doi=10.1038/nature.2012.10933 |s2cid=123219913 |issn=0028-0836 |access-date=January 1, 2019}}</ref> Direct fossil evidence of feathers or feather-like structures has been discovered in a diverse array of species in many non-avian dinosaur groups,<ref name="NYT-20161208" /> both among saurischians and ornithischians. Simple, branched, feather-like structures are known from ], primitive ]ns,<ref name="Godefroit2014">{{cite journal | last1 = Godefroit | first1 = P. | last2 = Sinitsa | first2 = S.M. | last3 = Dhouailly | first3 = D. | last4 = Bolotsky | first4 = Y.L. | last5 = Sizov | first5 = A.V. | last6 = McNamara | first6 = M.E. | last7 = Benton | first7 = M.J. | last8 = Spagna | first8 = P. | year = 2014 | title = A Jurassic ornithischian dinosaur from Siberia with both feathers and scales | url = http://palaeo.gly.bris.ac.uk/Benton/reprints/2014Kulinda.pdf | journal = Science | volume = 345 | issue = 6195 | pages = 451–455 | doi = 10.1126/science.1253351 | pmid = 25061209 | bibcode = 2014Sci...345..451G | hdl = 1983/a7ae6dfb-55bf-4ca4-bd8b-a5ea5f323103 | s2cid = 206556907 | access-date = July 27, 2016 | archive-url = https://web.archive.org/web/20190209232112/http://palaeo.gly.bris.ac.uk/Benton/reprints/2014Kulinda.pdf | archive-date = February 9, 2019 | url-status = dead }}</ref> and theropods,<ref name=Xuetal2004/> and primitive ceratopsians. Evidence for true, vaned feathers similar to the flight feathers of modern birds has been found only in the theropod subgroup Maniraptora, which includes oviraptorosaurs, troodontids, dromaeosaurids, and birds.<ref name=KP04/><ref name=GC06/> Feather-like structures known as ]s have also been found in pterosaurs.<ref name=kellneretal2009>{{cite journal |last1=Kellner |first1=Alexander W. A. |author1-link=Alexander Kellner |last2=Wang |first2=Xiaolin |last3=Tischlinger |first3=Helmut |last4=Campos |first4=Diogenes de Almeida |last5=Hone |first5=David W. E. |last6=Meng |first6=Xi |display-authors=3 |year=2010 |title=The soft tissue of ''Jeholopterus'' (Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane |journal=Proceedings of the Royal Society B |location=London |publisher=Royal Society |volume=277 |issue=1679 |pages=321–329 |doi=10.1098/rspb.2009.0846 |issn=0962-8452 |pmc=2842671 |pmid=19656798}}</ref>


However, researchers do not agree regarding whether these structures share a common origin between lineages (i.e., they are ]),<ref name="mayr2016">{{cite journal |last1=Mayr |first1=G. |last2=Pittman |first2=M. |last3=Saitta |first3=E. |last4=Kaye |first4=T.G. |last5=Vinther |first5=J. |year=2016 |title=Structure and homology of ''Psittacosaurus'' tail bristles |journal=Palaeontology |volume=59 |issue=6 |pages=793–802 |doi=10.1111/pala.12257|bibcode=2016Palgy..59..793M |hdl=1983/029c668f-08b9-45f6-a0c5-30ce9256e593 |s2cid=89156313|url=https://research-information.bris.ac.uk/en/publications/029c668f-08b9-45f6-a0c5-30ce9256e593 |hdl-access=free }}</ref><ref name="benton2019"/> or if they were the result of widespread experimentation with skin coverings among ornithodirans.<ref name="barrett2015">{{cite journal |last1=Barrett |first1=P.M. |last2=Evans |first2=D.C. |last3=Campione |first3=N.E. |year=2015 |title=Evolution of dinosaur epidermal structures |journal=Biology Letters |volume=11 |issue=6 |pages=20150229 |doi=10.1098/rsbl.2015.0229 |pmc=4528472 |pmid=26041865}}</ref> If the former is the case, filaments may have been common in the ornithodiran lineage and evolved before the appearance of dinosaurs themselves.<ref name=switeknature/> Research into the genetics of ]s has revealed that crocodylian ]s do possess feather-keratins during embryonic development, but these keratins are not expressed by the animals before hatching.<ref name=AlibL2006>{{cite journal |last1=Alibardi |first1=Lorenzo |last2=Knapp |first2=Loren W. |last3=Sawyer |first3=Roger H. |year=2006 |title=Beta-keratin localization in developing alligator scales and feathers in relation to the development and evolution of feathers |journal=Journal of Submicroscopic Cytology and Pathology |location=] |publisher=Nuova Immagine Editrice |volume=38 |issue=2–3 |pages=175–192 |issn=1122-9497 |pmid=17784647}}</ref> The description of feathered dinosaurs has not been without controversy in general; perhaps the most vocal critics have been Alan Feduccia and Theagarten Lingham-Soliar, who have proposed that some purported feather-like fossils are the result of the decomposition of collagenous fiber that underlaid the dinosaurs' skin,<ref name=TLS03/><ref name=FLH05/><ref name=LSFX07/> and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with dinosaurs.<ref name=AF02/><ref name=FLH05/> However, their views have for the most part not been accepted by other researchers, to the point that the scientific nature of Feduccia's proposals has been questioned.<ref name=Prum2003/>
]
]
]


'']'' was the first fossil found that revealed a potential connection between dinosaurs and birds. It is considered a ], in that it displays features of both groups. Brought to light just two years after ]'s seminal '']'' (1859), its discovery spurred the nascent debate between proponents of ] and ]. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, at least one specimen was mistaken for the small theropod '']''.<ref name=PW88/> Since the 1990s, a number of additional feathered dinosaurs have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Many of these specimens were unearthed in the ] of the Jehol Biota.<ref name="benton2019">{{cite journal |last1=Benton |first1=M.J. |last2=Dhouailly |first2=D. |last3=Jiang |first3=B. |last4=McNamara |first4=M. |year=2019 |title=The Early Origin of Feathers |journal=Trends in Ecology & Evolution |volume=34 |issue=9 |pages=856–869 |doi=10.1016/j.tree.2019.04.018|pmid=31164250 |bibcode=2019TEcoE..34..856B |hdl=10468/8068 |s2cid=174811556 |hdl-access=free }}</ref> If feather-like structures were indeed widely present among non-avian dinosaurs, the lack of abundant fossil evidence for them may be due to the fact that delicate features like skin and feathers are seldom preserved by fossilization and thus often absent from the fossil record.<ref name="Schweitzeretal1999">{{cite journal |last1=Schweitzer |first1=Mary H. |last2=Watt |first2=J.A. |last3=Avci |first3=R. |last4=Knapp |first4=L. |last5=Chiappe |first5=L. |last6=Norell |first6=M. |last7=Marshall |first7=M. |display-authors=3 |year=1999 |title=Beta-keratin specific immunological reactivity in feather-like structures of the Cretaceous Alvarezsaurid, ''Shuvuuia deserti'' |journal=] |location=Hoboken, NJ |publisher=Wiley-Blackwell |volume=285 |issue=2 |pages=146–157 |pmid=10440726 |doi=10.1002/(SICI)1097-010X(19990815)285:2<146::AID-JEZ7>3.0.CO;2-A |bibcode=1999JEZ...285..146S |issn=1552-5007}}</ref>
{{Link FA|sl}}
{{Link FA|th}}
{{Link FA|es}}


===Skeleton===
]
Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent another important line of evidence for paleontologists. Areas of the skeleton with important similarities include the neck, pubis, ] (semi-lunate ]), arm and ], furcula (wishbone), and ]. Comparison of bird and dinosaur skeletons through cladistic analysis strengthens the case for the link.<ref name=archaeopteryxucmp>{{cite web |url=https://ucmp.berkeley.edu/diapsids/birds/archaeopteryx.html |last=<!--Staff writer(s); no by-line.--> |title=''Archaeopteryx'': An Early Bird |publisher=University of California Museum of Paleontology |location=Berkeley |access-date=October 30, 2019}}</ref>
]

]
===Soft anatomy===
]
] of '']'']]
]
Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to a 2005 investigation led by Patrick M. O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said.<ref name=OConnorClaessens2005/><ref>{{Cite journal |last=O'Connor |first=Patrick M. |last2=Claessens |first2=Leon P. A. M. |date=July 2005 |title=Basic avian pulmonary design and flow-through ventilation in non-avian theropod dinosaurs |url=https://www.nature.com/articles/nature03716 |journal=Nature |language=en |volume=436 |issue=7048 |pages=253–256 |doi=10.1038/nature03716 |issn=0028-0836}}</ref> In 2008, scientists described '']'', the skeleton of which supplies the strongest evidence to date of a dinosaur with a bird-like breathing system. ]ning of ''Aerosteon'''s fossil bones revealed evidence for the existence of air sacs within the animal's body cavity.<ref name=Sereno2008/><ref name=newswise2/>
]

]
===Behavioral evidence===
]
Fossils of the troodonts '']'' and '']'' demonstrate that some dinosaurs slept with their heads tucked under their arms.<ref name=XUNorell2004/> This behavior, which may have helped to keep the head warm, is also characteristic of modern birds. Several ] and oviraptorosaur specimens have also been found preserved on top of their nests, likely brooding in a bird-like manner.<ref name="norell1995">{{cite journal |last1=Norell |first1=Mark A. |last2=Clark |first2=James M. |last3=Chiappe |first3=Luis M. |last4=Dashzeveg |first4=Demberelyin |display-authors=3 |year=1995 |title=A nesting dinosaur |journal=Nature |location=London |publisher=Nature Research |volume=378 |issue=6559 |pages=774–776 |bibcode=1995Natur.378..774N |doi=10.1038/378774a0 |s2cid=4245228 |issn=0028-0836}}</ref> The ratio between egg volume and body mass of adults among these dinosaurs suggest that the eggs were primarily brooded by the male and that the young were highly precocial, similar to many modern ground-dwelling birds.<ref name=Varricchioetal2008>{{cite journal |last1=Varricchio |first1=David J. |last2=Moore |first2=Jason R. |last3=Erickson |first3=Gregory M. |author3-link=Gregory M. Erickson |last4=Norell |first4=Mark A. |last5=Jackson |first5=Frankie D. |last6=Borkowski |first6=John J. |display-authors=3 |year=2008 |title=Avian Paternal Care Had Dinosaur Origin |journal=Science |location=Washington, D.C. |publisher=American Association for the Advancement of Science |volume=322 |issue=5909 |pages=1826–1828 |bibcode=2008Sci...322.1826V |doi=10.1126/science.1163245 |issn=0036-8075 |pmid=19095938|s2cid=8718747 |doi-access=free }}</ref>
]

]
Some dinosaurs are known to have used ] stones like modern birds. These stones are swallowed by animals to aid digestion and break down food and hard fibers once they enter the stomach. When found in association with fossils, gizzard stones are called gastroliths.<ref name=wings2007/>
]

]
==Extinction of major groups==
]
{{Main|Cretaceous–Paleogene extinction event}}
]
All non-avian dinosaurs and most lineages of birds<ref name="longrich2011">{{cite journal |last1=Longrich |first1=N.R. |last2=Tokaryk |first2=T. |last3=Field |first3=D.J. |year=2011 |title=Mass extinction of birds at the Cretaceous–Paleogene (K–Pg) boundary |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=37 |pages=15253–15257 |doi=10.1073/pnas.1110395108|pmid=21914849 |pmc=3174646 |bibcode=2011PNAS..10815253L |doi-access=free }}</ref> became extinct in a ], called the ], at the end of the Cretaceous period. Above the ], which has been dated to 66.038 ± 0.025 million years ago,<ref name="renne2013">{{cite journal |last1=Renne |first1=P.R. |last2=Deino |first2=A.L. |last3=Hilgen |first3=F.J. |last4=Kuiper |first4=K.F. |last5=Mark |first5=D.F. |last6=Mitchell |first6=W.S. |last7=Morgan |first7=L.E. |last8=Mundil |first8=R. |last9=Smit |first9=J. |year=2013 |title=Time scales of critical events around the Cretaceous-Paleogene boundary |journal=Science |volume=339 |issue=6120 |pages=684–687 |doi=10.1126/science.1230492|pmid=23393261 |bibcode=2013Sci...339..684R |s2cid=6112274 }}</ref> fossils of non-avian dinosaurs disappear abruptly; the absence of dinosaur fossils was historically used to assign rocks to the ensuing Cenozoic. The nature of the event that caused this mass extinction has been extensively studied since the 1970s, leading to the development of two mechanisms that are thought to have played major roles: an extraterrestrial ] in the ], along with ] volcanism in ]. However, the specific mechanisms of the extinction event and the extent of its effects on dinosaurs are still areas of ongoing research.<ref name="brusatte2014">{{cite journal |last1=Brusatte |first1=S.L. |last2=Butler |first2=R.J. |last3=Barrett |first3=P.M. |last4=Carrano |first4=M.T. |last5=Evans |first5=D.C. |last6=Lloyd |first6=G.T. |last7=Mannion |first7=P.D. |last8=Norell |first8=M.A. |last9=Peppe |first9=D.J. |last10=Upchurch |first10=P. |last11=Williamson |first11=T.E. |year=2015 |title=The extinction of the dinosaurs |journal=Biological Reviews |volume=90 |issue=2 |pages=628–642 |doi=10.1111/brv.12128|pmid=25065505 |hdl=20.500.11820/176e5907-26ec-4959-867f-0f2e52335f88 |s2cid=115134484 |url=https://www.research.ed.ac.uk/en/publications/176e5907-26ec-4959-867f-0f2e52335f88 |doi-access=free |hdl-access=free }}</ref> Alongside dinosaurs, many other groups of animals became extinct: pterosaurs, marine reptiles such as mosasaurs and plesiosaurs, several groups of mammals, ] (]-like ]), ]s (]-building ]), and various groups of marine plankton.<ref name="macleod1997"/><ref name="archibald1982"/> In all, approximately 47% of genera and 76% of species on Earth became extinct during the K-Pg extinction event.<ref name="jablonski1991">{{cite journal |last1=Jablonski |first1=D. |year=1991 |title=Extinctions: a paleontological perspective |journal=Science |volume=253 |issue=5021 |pages=754–757 |doi=10.1126/science.253.5021.754|pmid=17835491 |bibcode=1991Sci...253..754J }}</ref> The relatively large size of most dinosaurs and the low diversity of small-bodied dinosaur species at the end of the Cretaceous may have contributed to their extinction;<ref>{{cite journal |last1=Longrich |first1=N.R. |last2=Bhullar |first2=B.-A. S. |last3=Gauthier |first3=J.A. |year=2012 |title=Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary |journal=Proceedings of the National Academy of Sciences |volume=109 |issue=52 |pages=21396–21401 |bibcode=2012PNAS..10921396L |doi=10.1073/pnas.1211526110 |issn=0027-8424 |pmc=3535637 |pmid=23236177|doi-access=free }}</ref> the extinction of the bird lineages that did not survive may also have been caused by a dependence on forest habitats or a lack of adaptations to ] for survival.<ref name="field2018">{{cite journal |last1=Field |first1=D.J. |last2=Bercovici |first2=A. |last3=Berv |first3=J.S. |last4=Dunn |first4=R. |last5=Fastovsky |first5=D.E. |last6=Lyson |first6=T.R. |last7=Vajda |first7=V. |last8=Gauthier |first8=J.A. |year=2018 |title=Early evolution of modern birds structured by global forest collapse at the end-Cretaceous mass extinction |journal=Current Biology |volume=28 |issue=11 |pages=1825–1831 |doi=10.1016/j.cub.2018.04.062|pmid=29804807 |s2cid=44075214 |doi-access=free |bibcode=2018CBio...28E1825F }}</ref><ref name="larson2016">{{cite journal |last1=Larson |first1=D.W. |last2=Brown |first2=C.M. |last3=Evans |first3=D.C. |year=2016 |title=Dental disparity and ecological stability in bird-like dinosaurs prior to the end-Cretaceous mass extinction |journal=Current Biology |volume=26 |issue=10 |pages=1325–1333 |doi=10.1016/j.cub.2016.03.039|pmid=27112293 |s2cid=3937001 |doi-access=free |bibcode=2016CBio...26.1325L }}</ref>
]

]
===Pre-extinction diversity===
]
Just before the K-Pg extinction event, the number of non-avian dinosaur species that existed globally has been estimated at between 628 and 1078.<ref name="leloeuff2012">{{cite journal |last=Le Loeuff |first=J. |year=2012 |title=Paleobiogeography and biodiversity of Late Maastrichtian dinosaurs: how many dinosaur species went extinct at the Cretaceous-Tertiary boundary? |journal=] |volume=183 |issue=6 |pages=547–559 |doi=10.2113/gssgfbull.183.6.547 |issn=0037-9409}}</ref> It remains uncertain whether the diversity of dinosaurs was in gradual decline before the K-Pg extinction event, or whether dinosaurs were actually thriving prior to the extinction. Rock formations from the ] epoch, which directly preceded the extinction, have been found to have lower diversity than the preceding ] epoch, which led to the prevailing view of a long-term decline in diversity.<ref name="macleod1997"/><ref name="archibald1982">{{cite journal |last1=Archibald |first1=J.D. |last2=Clemens |first2=W.A. |year=1982 |title=Late Cretaceous Extinctions |journal=American Scientist |volume=70 |issue=4 |pages=377–385 |jstor=27851545|bibcode=1982AmSci..70..377A }}</ref><ref name="carpenter1983">{{cite journal |last1=Carpenter |first1=K. |year=1983 |title=Evidence suggesting gradual extinction of latest Cretaceous dinosaurs |journal=Naturwissenschaften |volume=70 |issue=12 |pages=611–612 |doi=10.1007/BF00377404 |bibcode=1983NW.....70..611C |s2cid=20078285 }}</ref> However, these comparisons did not account either for varying ] between rock units or for different extents of exploration and excavation.<ref name="brusatte2014"/> In 1984, ] carried out an analysis to account for these biases, and found no evidence of a decline;<ref name="russell1984">{{cite journal |last1=Russell |first1=D.A. |year=1984 |title=The gradual decline of the dinosaurs—fact or fallacy? |journal=Nature |volume=307 |issue=5949 |pages=360–361 |doi=10.1038/307360a0|bibcode=1984Natur.307..360R |s2cid=4269426 }}</ref> another analysis by David Fastovsky and colleagues in 2004 even showed that dinosaur diversity continually increased until the extinction,<ref name="fastovsky2004">{{cite journal |last1=Fastovsky |first1=D.E. |last2=Huang |first2=Y. |last3=Hsu |first3=J. |last4=Martin-McNaughton |first4=J. |last5=Sheehan |first5=P.M. |last6=Weishampel |first6=D.B. |year=2004 |title=Shape of Mesozoic dinosaur richness |journal=Geology |volume=32 |issue=10 |pages=877–880 |doi=10.1130/G20695.1|bibcode=2004Geo....32..877F |url=http://doc.rero.ch/record/14984/files/PAL_E2134.pdf }}</ref> but this analysis has been rebutted.<ref name="sullivan2006">{{cite book |last1=Sullivan |first1=R.M. |year=2006 |chapter=The shape of Mesozoic dinosaur richness: a reassessment |editor-last1=Lucas |editor-first1=S.G. |editor-last2=Sullivan |editor-first2=R.M. |title=Late Cretaceous vertebrates from the Western Interior |series=New Mexico Museum of Natural History and Science Bulletin |volume=35 |pages=403–405 |chapter-url=https://books.google.com/books?id=rCDYCQAAQBAJ&pg=PA403}}</ref> Since then, different approaches based on statistics and mathematical models have variously supported either a sudden extinction<ref name="brusatte2014"/><ref name="leloeuff2012"/><ref name="chiarenza2019">{{cite journal |last1=Chiarenza |first1=A.A. |last2=Mannion |first2=P.D. |last3=Lunt |first3=D.J. |last4=Farnsworth |first4=A. |last5=Jones |first5=L.A. |last6=Kelland |first6=S.J. |last7=Allison |first7=P.A. |year=2019 |title=Ecological niche modelling does not support climatically-driven dinosaur diversity decline before the Cretaceous/Paleogene mass extinction |journal=Nature Communications |volume=10 |issue=1 |pages=1–14 |doi=10.1038/s41467-019-08997-2|pmid=30842410 |pmc=6403247 |bibcode=2019NatCo..10.1091C }}</ref> or a gradual decline.<ref name="lloyd2011">{{cite journal |last1=Lloyd |first1=G.T. |year=2012 |title=A refined modelling approach to assess the influence of sampling on palaeobiodiversity curves: new support for declining Cretaceous dinosaur richness |journal=Biology Letters |volume=8 |issue=1 |pages=123–126 |doi=10.1098/rsbl.2011.0210|pmid=21508029 |pmc=3259943 |s2cid=1376734 }}</ref><ref name="sakamoto2016">{{cite journal |last1=Sakamoto |first1=M. |last2=Benton |first2=M.J. |last3=Venditti |first3=C. |year=2016 |title=Dinosaurs in decline tens of millions of years before their final extinction |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=18 |pages=5036–5040 |doi=10.1073/pnas.1521478113|pmid=27092007 |pmc=4983840 |bibcode=2016PNAS..113.5036S |doi-access=free }}</ref> End-Cretaceous trends in diversity may have varied between dinosaur lineages: it has been suggested that sauropods were not in decline, while ornithischians and theropods were in decline.<ref name="barrett2009">{{cite journal |last1=Barrett |first1=P.M. |last2=McGowan |first2=A.J. |last3=Page |first3=V. |year=2009 |title=Dinosaur diversity and the rock record |journal=Proceedings of the Royal Society B: Biological Sciences |volume=276 |issue=1667 |pages=2667–2674 |doi=10.1098/rspb.2009.0352|pmid=19403535 |pmc=2686664 }}</ref><ref name="upchurch2011">{{cite journal |last1=Upchurch |first1=P. |last2=Mannion |first2=P.D. |last3=Benson |first3=R.B. |last4=Butler |first4=R.J. |last5=Carrano |first5=M.T. |year=2011 |title=Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria |journal=Geological Society, London, Special Publications |volume=358 |issue=1 |pages=209–240 |doi=10.1144/SP358.14|bibcode=2011GSLSP.358..209U |s2cid=130777837 }}</ref>
]

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===Impact event===
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{{Main|Chicxulub crater}}
]
] (left) and his son ] (right) at the K-T Boundary in ], Italy, 1981]]
]
] at the tip of the ]; the impactor that formed this crater may have caused the dinosaur ].]]
]
The ], first brought to wide attention in 1980 by ], ], and colleagues, attributes the K-Pg extinction event to a ] (extraterrestrial projectile) impact.<ref>{{harvnb|Randall|2015}}</ref> Alvarez and colleagues proposed that a sudden increase in ] levels, recorded around the world in rock deposits at the Cretaceous–Paleogene boundary, was direct evidence of the impact.<ref name="alvarez1980"/> ], indicative of a strong shockwave emanating from an impact, was also found worldwide.<ref name="bohor1987">{{cite journal |last1=Bohor |first1=B.F. |last2=Modreski |first2=P.J. |last3=Foord |first3=E.E. |year=1987 |title=Shocked quartz in the Cretaceous-Tertiary boundary clays: Evidence for a global distribution |journal=Science |volume=236 |issue=4802 |pages=705–709 |doi=10.1126/science.236.4802.705|pmid=17748309 |bibcode=1987Sci...236..705B |s2cid=31383614 |url=https://zenodo.org/record/1230978 }}</ref> The actual impact site remained elusive until a ] measuring {{convert|180|km|mi|abbr=on}} wide was discovered in the Yucatán Peninsula of southeastern ], and was publicized in a 1991 paper by ] and colleagues.<ref name="hildebrand1991">{{cite journal |last1=Hildebrand |first1=A.R. |last2=Penfield |first2=G.T. |last3=Kring |first3=D.A. |last4=Pilkington |first4=M. |last5=Camargo |first5=Z.A. |last6=Jacobsen |first6=S.B. |last7=Boynton |first7=W.V. |year=1991 |title=Chicxulub crater: a possible Cretaceous/Tertiary boundary impact crater on the Yucatan Peninsula, Mexico |journal=Geology |volume=19 |issue=9 |pages=867–871 |doi=10.1130/0091-7613(1991)019<0867:CCAPCT>2.3.CO;2|bibcode=1991Geo....19..867H }}</ref> Now, the bulk of the evidence suggests that a bolide {{convert|5|to|15|km|mi|frac=2|abbr=off|sp=us}} wide impacted the Yucatán Peninsula 66 million years ago, forming this crater<ref name="pope1996"/> and creating a "kill mechanism" that triggered the extinction event.<ref name="schulte2010">{{cite journal |last1=Schulte |first1=P. |last2=Alegret |first2=L. |last3=Arenillas |first3=I. |last4=Arz |first4=J.A. |last5=Barton |first5=P.J. |last6=Bown |first6=P.R. |last7=Bralower |first7=T.J. |last8=Christeson |first8=G.L. |last9=Claeys |first9=P. |last10=Cockell |first10=C.S. |last11=Collins |first11=G.S. |first12=A. |last12=Deutsch |first13=T.J. |last13=Goldin |first14=K. |last14=Goto |first15=J.M. |last15=Grajales-Nishimura |first16=R.A.F. |last16=Grieve |first17=S.P.S. |last17=Gulick |first18=K.R. |last18=Johnson |first19=W. |last19=Kiessling |first20=C. |last20=Koeberl |first21=D.A. |last21=Kring |first22=K.G. |last22=MacLeod |first23=T. |last23=Matsui |first24=J. |last24=Melosh |first25=A. |last25=Montanari |first26=J.V. |last26=Morgan |first27=C.R. |last27=Neal |first28=D.J. |last28=Nichols |first29=R.D. |last29=Norris |first30=E. |last30=Pierazzo |first31=G. |last31=Ravizza |first32=M. |last32=Rebolledo-Vieyra |first33=W. |last33=Uwe Reimold |first34=E. |last34=Robin |first35=T. |last35=Salge |first36=R.P. |last36=Speijer |first37=A.R. |last37=Sweet |first38=J. |last38=Urrutia-Fucugauchi |first39=V. |last39=Vajda |first40=M.T. |last40=Whalen |first41=P.S. |last41=Willumsen |year=2010 |title=The Chicxulub asteroid impact and mass extinction at the Cretaceous-Paleogene boundary |journal=Science |volume=327 |issue=5970 |pages=1214–1218 |doi=10.1126/science.1177265|pmid=20203042 |bibcode=2010Sci...327.1214S |s2cid=2659741 |url=https://lirias.kuleuven.be/handle/123456789/264213 }}</ref><ref name="kring2007">{{cite journal |last1=Kring |first1=D. A. |year=2007 |title=The Chicxulub impact event and its environmental consequences at the Cretaceous–Tertiary boundary |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=255 |issue=1–2 |pages=4–21 |doi=10.1016/j.palaeo.2007.02.037|bibcode=2007PPP...255....4K }}</ref><ref name="chiarenza2020">{{cite journal |last1=Chiarenza |first1=A.A. |last2=Farnsworth |first2=A. |last3=Mannion |first3=P.D. |last4=Lunt |first4=D.J. |last5=Valdes |first5=P.J. |last6=Morgan |first6=J.V.|author6-link= Joanna Morgan |last7=Allison |first7=P.A. |year=2020 |title=Asteroid impact, not volcanism, caused the end-Cretaceous dinosaur extinction |journal=Proceedings of the National Academy of Sciences |volume=117 |issue=29 |pages=17084–17093 |doi=10.1073/pnas.2006087117|pmid=32601204 |pmc=7382232 |bibcode=2020PNAS..11717084C |doi-access=free }}</ref>
]

]
Within hours, the Chicxulub impact would have created immediate effects such as earthquakes,<ref name="ivanov2005">{{cite journal |last1=Ivanov |first1=B.A. |year=2005 |title=Numerical Modeling of the Largest Terrestrial Meteorite Craters |journal=Solar System Research |volume=39 |issue=5 |pages=381–409 |doi=10.1007/s11208-005-0051-0|bibcode=2005SoSyR..39..381I |s2cid=120305483 }}</ref> tsunamis,<ref name="matsui2002">{{cite journal |last1=Matsui |first1=T. |last2=Imamura |first2=F. |last3=Tajika |first3=E. |last4=Nakano |first4=Y. |last5=Fujisawa |first5=Y. |year=2002 |title=Generation and propagation of a tsunami from the Cretaceous-Tertiary impact event |journal=Geological Society of America Special Papers |volume=356 |pages=69–78 |doi=10.1130/0-8137-2356-6.69|isbn=978-0-8137-2356-3 }}</ref> and a global firestorm that likely killed unsheltered animals and started wildfires.<ref name="robertson2004">{{cite journal |last1=Robertson |first1=D.S. |last2=McKenna |first2=M.C. |author2-link=Malcolm McKenna |last3=Toon |first3=O.B. |author3-link=Owen Toon |last4=Hope |first4=S. |last5=Lillegraven |first5=J.A. |display-authors=3 |year=2004 |title=Survival in the first hours of the Cenozoic |url=http://webh01.ua.ac.be/funmorph/raoul/macroevolutie/Robertson2004.pdf |url-status=dead |journal=] |volume=116 |issue=5–6 |pages=760–768 |bibcode=2004GSAB..116..760R |doi=10.1130/B25402.1 |issn=0016-7606 |archive-url=https://web.archive.org/web/20120918141759/http://webh01.ua.ac.be/funmorph/raoul/macroevolutie/Robertson2004.pdf |archive-date=September 18, 2012 |access-date=June 15, 2011}}</ref><ref name="robertson2013">{{cite journal |last1=Robertson |first1=D.S. |last2=Lewis |first2=W.M. |last3=Sheehan |first3=P.M. |last4=Toon |first4=O.B. |year=2013 |title=K-Pg extinction: Reevaluation of the heat-fire hypothesis |journal=Journal of Geophysical Research: Biogeosciences |volume=118 |issue=1 |pages=329–336 |doi=10.1002/jgrg.20018|bibcode=2013JGRG..118..329R |s2cid=17015462 |doi-access=free }}</ref> However, it would also have had longer-term consequences for the environment. Within days, sulfate ]s released from rocks at the impact site would have contributed to ] and ].<ref name="pope1997">{{cite journal |last1=Pope |first1=K.O. |last2=Baines |first2=K.H. |last3=Ocampo |first3=A.C. |last4=Ivanov |first4=B.A. |year=1997 |title=Energy, volatile production, and climatic effects of the Chicxulub Cretaceous/Tertiary impact |journal=Journal of Geophysical Research: Planets |volume=102 |issue=E9 |pages=21645–21664 |doi=10.1029/97JE01743|pmid=11541145 |bibcode=1997JGR...10221645P |s2cid=8447773 |doi-access=free }}</ref><ref name="ohno2014"/> ] aerosols are thought to have spread around the world over the ensuing months and years; they would have cooled the surface of the Earth by reflecting ], and greatly slowed ] by blocking out sunlight, thus creating an ].<ref name="brusatte2014"/><ref name="kaiho2016">{{cite journal |last1=Kaiho |first1=K. |last2=Oshima |first2=N. |last3=Adachi |first3=K. |last4=Adachi |first4=Y. |last5=Mizukami |first5=T. |last6=Fujibayashi |first6=M. |last7=Saito |first7=R. |year=2016 |title=Global climate change driven by soot at the K-Pg boundary as the cause of the mass extinction |journal=Scientific Reports |volume=6 |issue=1 |pages=1–13 |doi=10.1038/srep28427|pmid=27414998 |pmc=4944614 |bibcode=2016NatSR...628427K }}</ref><ref name="lyons2020">{{cite journal |last1=Lyons |first1=S.L. |last2=Karp |first2=A.T. |last3=Bralower |first3=T.J. |last4=Grice |first4=K. |last5=Schaefer |first5=B. |last6=Gulick |first6=S.P. |last7=Morgan |first7=J.V.|author7-link= Joanna Morgan |last8=Freeman |first8=K.H. |year=2020 |title=Organic matter from the Chicxulub crater exacerbated the K–Pg impact winter |journal=Proceedings of the National Academy of Sciences |volume=117 |issue=41 |pages=25327–25334 |doi=10.1073/pnas.2004596117|pmid=32989138 |pmc=7568312 |bibcode=2020PNAS..11725327L |doi-access=free }}</ref> (This role was ascribed to ] aerosols until experiments demonstrated otherwise.<ref name="ohno2014">{{cite journal |last1=Ohno |first1=S. |last2=Kadono |first2=T. |last3=Kurosawa |first3=K. |last4=Hamura |first4=T. |last5=Sakaiya |first5=T. |last6=Shigemori |first6=K. |last7=Hironaka |first7=Y. |last8=Sano |first8=T. |last9=Watari |first9=T. |last10=Otani |first10=K. |last11=Matsui |first11=T. |last12=Sugita |first12=S. |year=2014 |title=Production of sulphate-rich vapour during the Chicxulub impact and implications for ocean acidification |journal=Nature Geoscience |volume=7 |issue=4 |pages=279–282 |doi=10.1038/ngeo2095|bibcode=2014NatGe...7..279O }}</ref>) The cessation of photosynthesis would have led to the collapse of ]s depending on leafy plants, which included all dinosaurs save for grain-eating birds.<ref name="larson2016"/>
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===Deccan Traps===
]
{{Main|Deccan Traps}} <!-- This section is linked from ] -->
]

]
At the time of the K-Pg extinction, the ] flood basalts of India were actively erupting. The eruptions can be separated into three phases around the K-Pg boundary, two prior to the boundary and one after. The second phase, which occurred very close to the boundary, would have extruded 70 to 80% of the volume of these eruptions in intermittent pulses that occurred around 100,000 years apart.<ref name="chenet2009">{{cite journal |last1=Chenet |first1=A.L. |last2=Courtillot |first2=V. |last3=Fluteau |first3=F. |last4=Gérard |first4=M. |last5=Quidelleur |first5=X. |last6=Khadri |first6=S.F.R. |last7=Subbarao |first7=K.V. |last8=Thordarson |first8=T. |year=2009 |title=Determination of rapid Deccan eruptions across the Cretaceous-Tertiary boundary using paleomagnetic secular variation: 2. Constraints from analysis of eight new sections and synthesis for a 3500-m-thick composite section |journal=Journal of Geophysical Research: Solid Earth |volume=114 |issue=B6 |pages=B06103 |doi=10.1029/2008JB005644|bibcode=2009JGRB..114.6103C |s2cid=140541003 |url=https://hal.archives-ouvertes.fr/hal-00406544/file/2008JB005644.pdf }}</ref><ref name="schoene2019">{{cite journal |last1=Schoene |first1=B. |last2=Eddy |first2=M.P. |last3=Samperton |first3=K.M. |last4=Keller |first4=C.B. |last5=Keller |first5=G. |last6=Adatte |first6=T. |last7=Khadri |first7=S.F. |year=2019 |title=U-Pb constraints on pulsed eruption of the Deccan Traps across the end-Cretaceous mass extinction |journal=Science |volume=363 |issue=6429 |pages=862–866 |doi=10.1126/science.aau2422|pmid=30792300 |bibcode=2019Sci...363..862S |osti=1497969 |s2cid=67876950 |doi-access=free }}</ref> ] such as ] and ] would have been released by this volcanic activity,<ref name="mclean1985">{{cite journal |last1=McLean |first1=D.M. |year=1985 |title=Deccan Traps mantle degassing in the terminal Cretaceous marine extinctions |journal=Cretaceous Research |volume=6 |issue=3 |pages=235–259 |doi=10.1016/0195-6671(85)90048-5|bibcode=1985CrRes...6..235M }}</ref><ref name="self2006">{{cite journal |last1=Self |first1=S. |last2=Widdowson |first2=M. |last3=Thordarson |first3=T. |last4=Jay |first4=A.E. |year=2006 |title=Volatile fluxes during flood basalt eruptions and potential effects on the global environment: A Deccan perspective |journal=Earth and Planetary Science Letters |volume=248 |issue=1–2 |pages=518–532 |doi=10.1016/j.epsl.2006.05.041|bibcode=2006E&PSL.248..518S }}</ref> resulting in ] through temperature perturbations of roughly {{convert|3|C-change}} but possibly as high as {{convert|7|C-change}}.<ref name="tobin2017">{{cite journal |last1=Tobin |first1=T.S. |last2=Bitz |first2=C.M. |last3=Archer |first3=D. |year=2017 |title=Modeling climatic effects of carbon dioxide emissions from Deccan Traps volcanic eruptions around the Cretaceous–Paleogene boundary |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=478 |pages=139–148 |doi=10.1016/j.palaeo.2016.05.028|bibcode=2017PPP...478..139T }}</ref> Like the Chicxulub impact, the eruptions may also have released sulfate aerosols, which would have caused acid rain and global cooling.<ref name="schmidt2016">{{cite journal |last1=Schmidt |first1=A. |last2=Skeffington |first2=R.A. |last3=Thordarson |first3=T. |last4=Self |first4=S. |last5=Forster |first5=P.M. |last6=Rap |first6=A. |last7=Ridgwell |first7=A. |last8=Fowler |first8=D. |last9=Wilson |first9=M. |last10=Mann |first10=G.W. |last11=Wignall |first11=P.B. |last12=Carslaw |first12=K.S. |year=2016 |title=Selective environmental stress from sulphur emitted by continental flood basalt eruptions |journal=Nature Geoscience |volume=9 |issue=1 |pages=77–82 |doi=10.1038/ngeo2588|bibcode=2016NatGe...9...77S |s2cid=59518452 |url=http://eprints.whiterose.ac.uk/92239/1/Schmidt_et_al_NGS_accepted.pdf |archive-url=https://web.archive.org/web/20170922012438/http://eprints.whiterose.ac.uk/92239/1/Schmidt_et_al_NGS_accepted.pdf |archive-date=2017-09-22 |url-status=live }}</ref> However, due to large error margins in the dating of the eruptions, the role of the Deccan Traps in the K-Pg extinction remains unclear.<ref name="renne2013"/><ref name="brusatte2014"/><ref name="hofman2000"/>
]

]
Before 2000, arguments that the Deccan Traps eruptions—as opposed to the Chicxulub impact—caused the extinction were usually linked to the view that the extinction was gradual. Prior to the discovery of the Chicxulub crater, the Deccan Traps were used to explain the global iridium layer;<ref name="mclean1985"/><ref name="sahni1988">{{cite journal |last1=Sahni |first1=A. |year=1988 |title=Cretaceous-Tertiary boundary events: Mass extinctions, iridium enrichment and Deccan volcanism |journal=Current Science |volume=57 |issue=10 |pages=513–519 |jstor=24090754}}</ref> even after the crater's discovery, the impact was still thought to only have had a regional, not global, effect on the extinction event.<ref name="glasby1996">{{cite journal |last1=Glasby |first1=G.P. |last2=Kunzendorf |first2=H. |year=1996 |title=Multiple factors in the origin of the Cretaceous/Tertiary boundary: the role of environmental stress and Deccan Trap volcanism |journal=Geologische Rundschau |volume=85 |issue=2 |pages=191–210 |doi=10.1007/BF02422228|pmid=11543126 |bibcode=1996IJEaS..85..191G |s2cid=19155384 }}</ref> In response, Luis Alvarez rejected volcanic activity as an explanation for the iridium layer and the extinction as a whole.<ref name="alvarez1987">{{cite report |last1=Alvarez |first1=L.W. |year=1987 |title=Mass Extinctions Caused by Large Bolide Impacts |url=https://www.osti.gov/servlets/purl/875729 |docket=LBL-22786 |page=39 |publisher=] |access-date=January 27, 2021}}</ref> Since then, however, most researchers have adopted a more moderate position, which identifies the Chicxulub impact as the primary progenitor of the extinction while also recognizing that the Deccan Traps may also have played a role. Walter Alvarez himself has acknowledged that the Deccan Traps and other ecological factors may have contributed to the extinctions in addition to the Chicxulub impact.<ref name="alvarez1997"/> Some estimates have placed the start of the second phase in the Deccan Traps eruptions within 50,000 years after the Chicxulub impact.<ref name="renne2015">{{cite journal |last1=Renne |first1=P.R. |last2=Sprain |first2=C.J. |last3=Richards |first3=M.A. |last4=Self |first4=S. |last5=Vanderkluysen |first5=L. |last6=Pande |first6=K. |year=2015 |title=State shift in Deccan volcanism at the Cretaceous-Paleogene boundary, possibly induced by impact |journal=Science |volume=350 |issue=6256 |pages=76–78 |doi=10.1126/science.aac7549|pmid=26430116 |bibcode=2015Sci...350...76R |s2cid=30612906 |url=https://escholarship.org/uc/item/5fk799n8 |doi-access=free }}</ref> Combined with mathematical modelling of the ]s that would have been generated by the impact, this has led to the suggestion that the Chicxulub impact may have triggered these eruptions by increasing the permeability of the ] underlying the Deccan Traps.<ref name="richards2015">{{cite journal |last1=Richards |first1=M.A. |last2=Alvarez |first2=W. |last3=Self |first3=S. |last4=Karlstrom |first4=L. |last5=Renne |first5=P.R. |last6=Manga |first6=M. |last7=Sprain |first7=C.J. |last8=Smit |first8=J. |last9=Vanderkluysen |first9=L. |last10=Gibson |first10=S.A. |year=2015 |title=Triggering of the largest Deccan eruptions by the Chicxulub impact |journal=Geological Society of America Bulletin |volume=127 |issue=11–12 |pages=1507–1520 |doi=10.1130/B31167.1|bibcode=2015GSAB..127.1507R |s2cid=3463018 |url=https://escholarship.org/uc/item/86f3521g }}</ref><ref name="khazins2019">{{cite book |last1=Khazins |first1=V. |last2=Shuvalov |first2=V. |year=2019 |editor-first1=G. |editor-last1=Kocharyan |editor-first2=A. |editor-last2=Lyakhov |chapter=Chicxulub Impact as a Trigger of One of Deccan Volcanism Phases: Threshold of Seismic Energy Density |title=Trigger Effects in Geosystems |pages=523–530 |publisher=Springer |location=Cham |series=Springer Proceedings in Earth and Environmental Sciences |doi=10.1007/978-3-030-31970-0_55|isbn=978-3-030-31969-4 |s2cid=210277965 }}</ref>
]

]
Whether the Deccan Traps were a major cause of the extinction, on par with the Chicxulub impact, remains uncertain. Proponents consider the climatic impact of the sulfur dioxide released to have been on par with the Chicxulub impact, and also note the role of flood basalt volcanism in other mass extinctions like the ].<ref name="archibald2010">{{cite journal |first1=J.D. |last1=Archibald |first2=W.A. |last2=Clemens |first3=K. |last3=Padian |first4=T. |last4=Rowe |first5=N. |last5=Macleod |first6=P.M. |last6=Barrett |first7=A. |last7=Gale |first8=P. |last8=Holroyd |first9=H.-D. |last9=Sues |first10=N.C. |last10=Arens |first11=J.R. |last11=Horner |first12=G.P. |last12=Wilson |first13=M.B. |last13=Goodwin |first14=C.A. |last14=Brochu |first15=D.L. |last15=Lofgren |first16=S.H. |last16=Hurlbert |first17=J.H. |last17=Hartman |first18=D.A. |last18=Eberth |first19=P.B. |last19=Wignall |first20=P.J. |last20=Currie |first21=A. |last21=Weil |first22=G.V.R. |last22=Prasad |first23=L. |last23=Dingus |first24=V. |last24=Courtillot |first25=A. |last25=Milner |first26=A. |last26=Milner |first27=S. |last27=Bajpai |first28=D.J. |last28=Ward |first29=A. |last29=Sahni |year=2010 |title=Cretaceous extinctions: multiple causes |journal=Science |volume=328 |issue=5981 |pages=973; author reply 975–6 |doi=10.1126/science.328.5981.973-a|pmid=20489004 }}</ref><ref name="courtillot2010">{{cite journal |last1=Courtillot |first1=V. |last2=Fluteau |first2=F. |year=2010 |title=Cretaceous extinctions: the volcanic hypothesis |journal=Science |volume=328 |issue=5981 |pages=973–974 |doi=10.1126/science.328.5981.973-b|pmid=20489003 }}</ref> They consider the Chicxulub impact to have worsened the ongoing climate change caused by the eruptions.<ref name="keller2014">{{cite journal |last1=Keller |first1=G. |year=2014 |title=Deccan volcanism, the Chicxulub impact, and the end-Cretaceous mass extinction: Coincidence? Cause and effect |journal=Geological Society of America Special Papers |volume=505 |pages=57–89 |doi=10.1130/2014.2505(03)|isbn=978-0-8137-2505-5 }}</ref> Meanwhile, detractors point out the sudden nature of the extinction and that other pulses in Deccan Traps activity of comparable magnitude did not appear to have caused extinctions. They also contend that the causes of different mass extinctions should be assessed separately.<ref name="schulte2010reply">{{cite journal |last1=Schulte |first1=P. |last2=Alegret |first2=L. |last3=Arenillas |first3=I. |last4=Arz |first4=J.A. |last5=Barton |first5=P.J. |last6=Bown |first6=P.R. |last7=Bralower |first7=T.J. |last8=Christeson |first8=G.L. |last9=Claeys |first9=P. |last10=Cockell |first10=C.S. |last11=Collins |first11=G.S. |first12=A. |last12=Deutsch |first13=T.J. |last13=Goldin |first14=K. |last14=Goto |first15=J.M. |last15=Grajales-Nishimura |first16=R.A.F. |last16=Grieve |first17=S.P.S. |last17=Gulick |first18=K.R. |last18=Johnson |first19=W. |last19=Kiessling |first20=C. |last20=Koeberl |first21=D.A. |last21=Kring |first22=K.G. |last22=MacLeod |first23=T. |last23=Matsui |first24=J. |last24=Melosh |first25=A. |last25=Montanari |first26=J.V. |last26=Morgan|author26-link= Joanna Morgan |first27=C.R. |last27=Neal |first28=D.J. |last28=Nichols |first29=R.D. |last29=Norris |first30=E. |last30=Pierazzo |first31=G. |last31=Ravizza |first32=M. |last32=Rebolledo-Vieyra |first33=W. |last33=Uwe Reimold |first34=E. |last34=Robin |first35=T. |last35=Salge |first36=R.P. |last36=Speijer |first37=A.R. |last37=Sweet |first38=J. |last38=Urrutia-Fucugauchi |first39=V. |last39=Vajda |first40=M.T. |last40=Whalen |first41=P.S. |last41=Willumsen |year=2010 |title=Response—Cretaceous extinctions |journal=Science |volume=328 |issue=5981 |pages=975–976 |doi=10.1126/science.328.5981.975|url=https://lirias.kuleuven.be/handle/123456789/269158 }}</ref> In 2020, Alfio Chiarenza and colleagues suggested that the Deccan Traps may even have had the opposite effect: they suggested that the long-term warming caused by its carbon dioxide emissions may have dampened the impact winter from the Chicxulub impact.<ref name="chiarenza2020"/>
]

]
===Possible Paleocene survivors===
]
Non-avian dinosaur remains have occasionally been found above the K-Pg boundary. In 2000, ] and colleagues reported the discovery of a single hadrosaur right femur in the ] of ], and described it as evidence of Paleocene dinosaurs. The rock unit in which the bone was discovered has been dated to the early ] epoch, approximately 64.8 million years ago.<ref name="fassett2011"/> If the bone was not ] by weathering action, it would provide evidence that some dinosaur populations survived at least half a million years into the Cenozoic.<ref name="fassett2009"/> Other evidence includes the presence of dinosaur remains in the Hell Creek Formation up to {{convert|1.3|m|ft|abbr=on}} above the Cretaceous–Paleogene boundary, representing 40,000 years of elapsed time. This has been used to support the view that the K-Pg extinction was gradual.<ref name="sloan1986"/> However, these supposed Paleocene dinosaurs are considered by many other researchers to be ], that is, washed out of their original locations and then reburied in younger sediments.<ref name="lucas2009">{{cite journal |last1=Lucas |first1=S.G. |last2=Sullivan |first2=R.M. |last3=Cather |first3=S.M. |last4=Jasinski |first4=S.E. |last5=Fowler |first5=D.W. |last6=Heckert |first6=A.B. |last7=Spielmann |first7=J.A. |last8=Hunt |first8=A.P. |year=2009 |title=No definitive evidence of Paleocene dinosaurs in the San Juan Basin |journal=Palaeontologia Electronica |volume=12 |issue=2 |page=8A}}</ref><ref name="renne2012">{{cite journal |last1=Renne |first1=P.R. |last2=Goodwin |first2=M.B. |year=2012 |title=Direct U-Pb dating of Cretaceous and Paleocene dinosaur bones, San Juan Basin, New Mexico: COMMENT |journal=Geology |volume=40 |issue=4 |pages=e259 |doi=10.1130/G32521C.1|bibcode=2012Geo....40E.259R |doi-access=free }}</ref><ref name="lofgren1990">{{cite journal |last1=Lofgren |first1=D.L. |last2=Hotton |first2=C.L. |last3=Runkel |first3=A.C. |year=1990 |title=Reworking of Cretaceous dinosaurs into Paleocene channel, deposits, upper Hell Creek Formation, Montana |journal=Geology |volume=18 |issue=9 |pages=874–877 |doi=10.1130/0091-7613(1990)018<0874:ROCDIP>2.3.CO;2|bibcode=1990Geo....18..874L }}</ref> The age estimates have also been considered unreliable.<ref name="koenig2012">{{cite journal |last1=Koenig |first1=A.E. |last2=Lucas |first2=S.G. |last3=Neymark |first3=L.A. |last4=Heckert |first4=A.B. |last5=Sullivan |first5=R.M. |last6=Jasinski |first6=S.E. |last7=Fowler |first7=D.W. |year=2012 |title=Direct U-Pb dating of Cretaceous and Paleocene dinosaur bones, San Juan Basin, New Mexico: COMMENT |journal=Geology |volume=40 |issue=4 |pages=e262 |doi=10.1130/G32154C.1|bibcode=2012Geo....40E.262K |doi-access=free }}</ref>
]

]
==Cultural depictions==
]
{{Main|Cultural depictions of dinosaurs}}
]
] for the ] in 1853]]
]
]'' (1914) by ], featuring the first animated dinosaur]]
]
By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured the popular imagination and become an enduring part of human culture. The entry of the word "dinosaur" into the common ] reflects the animals' cultural importance: in English, "dinosaur" is commonly used to describe anything that is impractically large, obsolete, or bound for extinction.<ref name="m-w"/>
]

]
Public enthusiasm for dinosaurs first developed in ] England, where in 1854, three decades after the first scientific descriptions of dinosaur remains, a menagerie of lifelike ] was unveiled in ]'s ]. The Crystal Palace dinosaurs proved so popular that a strong market in smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened at parks and ] around the world, ensuring that successive generations would be introduced to the animals in an immersive and exciting way.<ref name=torrens1993/> The enduring popularity of dinosaurs, in its turn, has resulted in significant public funding for dinosaur science, and has frequently spurred new discoveries. In the United States, for example, the competition between museums for public attention led directly to the Bone Wars of the 1880s and 1890s, during which a pair of feuding paleontologists made enormous scientific contributions.<ref name=breithaupt1997/>
]

]
The popular preoccupation with dinosaurs has ensured their appearance in ], ], and other ]. Beginning in 1852 with a passing mention in ]{{'}} '']'',<ref name=bleakhouse/> dinosaurs have been featured in large numbers of ]al works. ]'s 1864 novel '']'', ]'s 1912 book '']'', the 1914 animated film '']'' (featuring the first animated dinosaur), the iconic 1933 ] '']'', the 1954 '']'' and its many sequels, the best-selling 1990 novel '']'' by ] and its 1993 ] are just a few notable examples of dinosaur appearances in fiction. Authors of general-interest ] works about dinosaurs, including some prominent paleontologists, have often sought to use the animals as a way to educate readers about science in general. Dinosaurs are ubiquitous in ]; numerous ] have referenced dinosaurs in printed or televised advertisements, either in order to sell their own products or in order to characterize their rivals as slow-moving, dim-witted, or obsolete.<ref name=DFGlut1997/><ref>{{cite book |last1=Lee |first1=Newton |last2=Madej |first2=Krystina |title=Disney Stories |chapter=Early Animation: Gags and Situations |date=2012 |pages=17–24 |doi=10.1007/978-1-4614-2101-6_3|isbn=978-1-4614-2100-9 |s2cid=192335675 }}</ref>
]

]
==See also==
]
* ]
]
* ]
]
* ]
]
* ]
]
* ]
* ]
* ]
* ]
{{Clear}}

==Notes==
{{Reflist|group=note}}

==References==
{{Reflist|refs=
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<ref name=Holland1909>This was recognized not later than 1909: {{cite web |url=http://www.hmnh.org/library/diplodocus/holland1910.html |url-status=dead |title=Dr. W. J. Holland and the Sprawling Sauropods |last=Celeskey |first=Matt |year=2005 |website=The Hairy Museum of Natural History |archive-url=https://web.archive.org/web/20110612180650/http://www.hmnh.org/library/diplodocus/holland1910.html |archive-date=June 12, 2011 |access-date=October 18, 2019}}
* {{cite journal |last=Holland |first=William J. |author-link=William Jacob Holland |date=May 1910 |title=A Review of Some Recent Criticisms of the Restorations of Sauropod Dinosaurs Existing in the Museums of the United States, with Special Reference to that of ''Diplodocus Carnegiei'' in the Carnegie Museum |url=https://archive.org/details/jstor-2455581 |journal=] |publisher=] |volume=44 |issue=521 |pages=259–283 |issn=0003-0147 |access-date=October 18, 2019|doi=10.1086/279138 |s2cid=84424110 }}
* The arguments and many of the images are also presented in {{harvnb|Desmond|1975}}.</ref>
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{{Refend}}

==Further reading==

{{Library resources box
|onlinebooks=yes
|by=no
|lcheading= Dinosaurs
|label=Dinosaurs
}}
* {{cite web |title=Two New Species of Large Predatory Dinosaur With Crocodile-Like Skulls Discovered on Isle of Wight |website=SciTechDaily |date=September 29, 2021 |author=University of Southampton|url=https://scitechdaily.com/two-new-species-of-large-predatory-dinosaur-with-crocodile-like-skulls-discovered-on-isle-of-wight/}}
* {{cite journal|last1=Zhou |first1=Zhonghe |date=October 2004 |title=The origin and early evolution of birds: discoveries, disputes, and perspectives from fossil evidence |url=http://www.cisneros-heredia.org/infotrans/usfq/ornitofauna/pdfs/zhou2004.pdf |url-status=dead |journal=Naturwissenschaften |location=Berlin |publisher=Springer Science+Business Media |volume=91|issue=10 |pages=455–471 |bibcode=2004NW.....91..455Z |doi=10.1007/s00114-004-0570-4|issn=0028-1042 |pmid=15365634 |s2cid=3329625 |archive-url=https://web.archive.org/web/20110721144552/http://www.cisneros-heredia.org/infotrans/usfq/ornitofauna/pdfs/zhou2004.pdf |archive-date=July 21, 2011 |access-date=November 6, 2019}}
* {{cite book |last=Paul |first=Gregory S. |year=2002 |title=Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds |location=]; London |publisher=]|isbn=978-0-8018-6763-7 |lccn=2001000242 |oclc=1088130487 |url-access=registration |url=https://archive.org/details/dinosaursofairev0000paul}}.
* {{cite book |author=Stewart, Tabori & Chang |year=1997 |title=The Humongous Book of Dinosaurs |location=New York |publisher=] |isbn=978-1-55670-596-0 |lccn=97000398|oclc=1037269801 |title-link=The Humongous Book of Dinosaurs}}
* {{cite book |last=Sternberg |first=Charles Mortram |author-link=Charles Mortram Sternberg |year=1966 |orig-year=Original edition published by E. Cloutier, printer to the King, 1946 |title=Canadian Dinosaurs|series=Geological Series |volume=54|edition=2nd |location=] |publisher=] |lccn=gs46000214 |oclc=1032865683}}

{{Avemetatarsalia|state=autocollapse}}
{{Birds}}
{{Dinosaurs by Continent}}
{{subject bar|Dinosaurs|Paleontology|d=y|s=Portal:Dinosaurs|auto=1}}
{{Taxonbar|from=Q430}}
{{Authority control}}

{{DEFAULTSORT:Dinosaur}}
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Latest revision as of 05:10, 27 December 2024

Archosaurian reptiles that dominated the Mesozoic Era For other uses, see Dinosaur (disambiguation). Not to be confused with Dinosaurus.

Dinosaurs
Temporal range: Late TriassicPresent, 233.23 – 0 Mya (range includes birds) PreꞒ O S D C P T J K Pg N (possible Middle Triassic record)
Herrerasaurus ischigualastensis
(a carnivorous basal dinosaur)Triceratops horridus
(a ceratopsian)Stegosaurus stenops
(a stegosaur)Apatosaurus louisae
(a sauropod)Edmontosaurus regalis
(a hadrosaurid ornithopod)Microraptor gui
(a dromaeosaurid theropod)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Sauropsida
Clade: Archosauria
Clade: Avemetatarsalia
Clade: Ornithodira
Clade: Dinosauromorpha
Clade: Dinosauriformes
Clade: Dracohors
Clade: Dinosauria
Owen, 1842
Major groups
Dinosaurs and possible dinosaurs of uncertain affinity
montage of four birds
Birds are avian dinosaurs, and in phylogenetic taxonomy their over 11,000 extant species are included in the group Dinosauria.

Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is a subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 mya and their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record shows that birds are feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch, and are the only dinosaur lineage known to have survived the Cretaceous–Paleogene extinction event approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which are all dinosaurs other than birds.

Dinosaurs are varied from taxonomic, morphological and ecological standpoints. Birds, at over 11,000 living species, are among the most diverse groups of vertebrates. Using fossil evidence, paleontologists have identified over 900 distinct genera and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized as dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence suggests that all dinosaurs were egg-laying, and that nest-building was a trait shared by many dinosaurs, both avian and non-avian.

While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed skeletal modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest sauropod dinosaurs are estimated to have reached lengths of 39.7 meters (130 feet) and heights of 18 m (59 ft) and were the largest land animals of all time. The misconception that non-avian dinosaurs were uniformly gigantic is based in part on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small, some measuring about 50 centimeters (20 inches) in length.

The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" (meaning "terrible lizard") being coined by Sir Richard Owen in 1842 to refer to these "great fossil lizards". Since then, mounted fossil dinosaur skeletons have been major attractions at museums worldwide, and dinosaurs have become an enduring part of popular culture. The large sizes of some dinosaurs, as well as their seemingly monstrous and fantastic nature, have ensured their regular appearance in best-selling books and films, such as the Jurassic Park franchise. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Definition

Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of the most recent common ancestor (MRCA) of Triceratops and modern birds (Neornithes), and all its descendants. It has also been suggested that Dinosauria be defined with respect to the MRCA of Megalosaurus and Iguanodon, because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria. Both definitions cover the same known genera: Dinosauria = Ornithischia + Saurischia. This includes major groups such as ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous quadrupeds), ceratopsians (bipedal or quadrupedal herbivores with neck frills), pachycephalosaurians (bipedal herbivores with thick skulls), ornithopods (bipedal or quadrupedal herbivores including "duck-bills"), theropods (mostly bipedal carnivores and birds), and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and tails).

Birds are the sole surviving dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct superorder. However, most contemporary paleontologists reject the traditional style of classification based on anatomical similarity, in favor of phylogenetic taxonomy based on deduced ancestry, in which each group is defined as all descendants of a given founding genus. Birds belong to the dinosaur subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians.

Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in 2017 suggested a radical revision of dinosaurian systematics. Phylogenetic analysis by Baron et al. recovered the Ornithischia as being closer to the Theropoda than the Sauropodomorpha, as opposed to the traditional union of theropods with sauropodomorphs. This would cause sauropods and kin to fall outside traditional dinosaurs, so they re-defined Dinosauria as the last common ancestor of Triceratops horridus, Passer domesticus and Diplodocus carnegii, and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. They also resurrected the clade Ornithoscelida to refer to the group containing Ornithischia and Theropoda.

General description

Triceratops skeleton, Natural History Museum of Los Angeles County

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind limbs held erect beneath the body. Other prehistoric animals, including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs, and Dimetrodon, while often popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs, being members of the clade Ornithodira. The other groups mentioned are, like dinosaurs and pterosaurs, members of Sauropsida (the reptile and bird clade), except Dimetrodon (which is a synapsid). None of them had the erect hind limb posture characteristic of true dinosaurs.

Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic Era, especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; mammals, for example, rarely exceeded the size of a domestic cat and were generally rodent-sized carnivores of small prey. Dinosaurs have always been recognized as an extremely varied group: over 900 non-avian dinosaur genera have been confidently identified (2018) with 1124 species (2016). Estimates put the total number of dinosaur genera preserved in the fossil record at 1850, nearly 75% still undiscovered, and the number that ever existed (in or out of the fossil record) at 3,400. A 2016 estimate put the number of dinosaur species living in the Mesozoic at 1,543–2,468, compared to the number of modern-day birds (avian dinosaurs) at 10,806 species.

Extinct dinosaurs, as well as modern birds, include genera that are herbivorous and others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some evolved into quadrupeds, and others, such as Anchisaurus and Iguanodon, could walk as easily on two or four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony armor. Although the best-known genera are remarkable for their large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global distribution by the Early Jurassic epoch at latest. Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs (such as Microraptor) could fly or at least glide, and others, such as spinosaurids, had semiaquatic habits.

Distinguishing anatomical features

While recent discoveries have made it more difficult to present a universally agreed-upon list of their distinguishing features, nearly all dinosaurs discovered so far share certain modifications to the ancestral archosaurian skeleton, or are clearly descendants of older dinosaurs showing these modifications. Although some later groups of dinosaurs featured further modified versions of these traits, they are considered typical for Dinosauria; the earliest dinosaurs had them and passed them on to their descendants. Such modifications, originating in the most recent common ancestor of a certain taxonomic group, are called the synapomorphies of such a group.

Labeled diagram of a typical archosaur skull, the skull of Dromaeosaurus

A detailed assessment of archosaur interrelations by Sterling Nesbitt confirmed or found the following twelve unambiguous synapomorphies, some previously known:

  • In the skull, a supratemporal fossa (excavation) is present in front of the supratemporal fenestra, the main opening in the rear skull roof
  • Epipophyses, obliquely backward-pointing processes on the rear top corners of the anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae
  • Apex of a deltopectoral crest (a projection on which the deltopectoral muscles attach) located at or more than 30% down the length of the humerus (upper arm bone)
  • Radius, a lower arm bone, shorter than 80% of humerus length
  • Fourth trochanter (projection where the caudofemoralis muscle attaches on the inner rear shaft) on the femur (thigh bone) is a sharp flange
  • Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft
  • On the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the top connecting surface, for the fibula occupies less than 30% of the transverse width of the element
  • Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity, the inner space of the braincase
  • In the pelvis, the proximal articular surfaces of the ischium with the ilium and the pubis are separated by a large concave surface (on the upper side of the ischium a part of the open hip joint is located between the contacts with the pubic bone and the ilium)
  • Cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs anterolaterally (curves to the front and the outer side)
  • Distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia (the rear surface of the lower end of the shinbone)
  • Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it touches the fibula, has a hollow profile)

Nesbitt found a number of further potential synapomorphies and discounted a number of synapomorphies previously suggested. Some of these are also present in silesaurids, which Nesbitt recovered as a sister group to Dinosauria, including a large anterior trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and pubis, the presence of a cnemial crest on the tibia and of an ascending process on the astragalus, and many others.

Hip joints and hindlimb postures of: (left to right) typical reptiles (sprawling), dinosaurs and mammals (erect), and rauisuchians (pillar-erect)

A variety of other skeletal features are shared by dinosaurs. However, because they either are common to other groups of archosaurs or were not present in all early dinosaurs, these features are not considered to be synapomorphies. For example, as diapsids, dinosaurs ancestrally had two pairs of Infratemporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the snout and lower jaw. Additionally, several characteristics once thought to be synapomorphies are now known to have appeared before dinosaurs, or were absent in the earliest dinosaurs and independently evolved by different dinosaur groups. These include an elongated scapula, or shoulder blade; a sacrum composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in Herrerasaurus); and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface (closed in Saturnalia tupiniquim, for example). Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic epoch are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature.

Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side. This posture is due to the development of a laterally facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur. Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles. Erect limbs probably also helped support the evolution of large size by reducing bending stresses on limbs. Some non-dinosaurian archosaurs, including rauisuchians, also had erect limbs but achieved this by a "pillar-erect" configuration of the hip joint, where instead of having a projection from the femur insert on a socket on the hip, the upper pelvic bone was rotated to form an overhanging shelf.

History of study

Further information: History of paleontology

Pre-scientific history

Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese considered them to be dragon bones and documented them as such. For example, Huayang Guo Zhi (華陽國志), a gazetteer compiled by Chang Qu (常璩) during the Western Jin Dynasty (265–316), reported the discovery of dragon bones at Wucheng in Sichuan Province. Villagers in central China have long unearthed fossilized "dragon bones" for use in traditional medicines. In Europe, dinosaur fossils were generally believed to be the remains of giants and other biblical creatures.

Early dinosaur research

William Buckland

Scholarly descriptions of what would now be recognized as dinosaur bones first appeared in the late 17th century in England. Part of a bone, now known to have been the femur of a Megalosaurus, was recovered from a limestone quarry at Cornwell near Chipping Norton, Oxfordshire, in 1676. The fragment was sent to Robert Plot, Professor of Chemistry at the University of Oxford and first curator of the Ashmolean Museum, who published a description in his The Natural History of Oxford-shire (1677). He correctly identified the bone as the lower extremity of the femur of a large animal, and recognized that it was too large to belong to any known species. He therefore concluded it to be the femur of a huge human, perhaps a Titan or another type of giant featured in legends. Edward Lhuyd, a friend of Sir Isaac Newton, published Lithophylacii Britannici ichnographia (1699), the first scientific treatment of what would now be recognized as a dinosaur. In it he described and named a sauropod tooth, "Rutellum impicatum", that had been found in Caswell, near Witney, Oxfordshire.

Sir Richard Owen's coining of the word dinosaur, in the 1842 revised version of his talk at an 1841 meeting of the British Association for the Advancement of Science.

Between 1815 and 1824, the Rev William Buckland, the first Reader of Geology at the University of Oxford, collected more fossilized bones of Megalosaurus and became the first person to describe a non-avian dinosaur in a scientific journal. The second non-avian dinosaur genus to be identified, Iguanodon, was purportedly discovered in 1822 by Mary Ann Mantell, the wife of English geologist Gideon Mantell, though this is disputed and some historians say Gideon had acquired remains years earlier. Gideon Mantell recognized similarities between his fossils and the bones of modern iguanas and published his findings in 1825.

The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English paleontologist Sir Richard Owen coined the term "dinosaur", using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world. The term is derived from Ancient Greek δεινός (deinos) 'terrible, potent or fearfully great' and σαῦρος (sauros) 'lizard or reptile'. Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it also to evoke their size and majesty. Owen recognized that the remains that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus, shared distinctive features, and so decided to present them as a distinct taxonomic group. As clarified by British geologist and historian Hugh Torrens, Owen had given a presentation about fossil reptiles to the British Association for the Advancement of Science in 1841, but reports of the time show that Owen did not mention the word "dinosaur", nor recognize dinosaurs as a distinct group of reptiles in his address. He introduced the Dinosauria only in the revised text version of his talk published in April 1842. With the backing of Prince Albert, the husband of Queen Victoria, Owen established the Natural History Museum, London, to display the national collection of dinosaur fossils and other biological and geological exhibits.

Discoveries in North America

Edward Drinker CopeOthniel Charles Marsh

In 1858, William Parker Foulke discovered the first known American dinosaur, in marl pits in the small town of Haddonfield, New Jersey. (Although fossils had been found before, their nature had not been correctly discerned.) The creature was named Hadrosaurus foulkii. It was an extremely important find: Hadrosaurus was one of the first nearly complete dinosaur skeletons found (the first was in 1834, in Maidstone, England), and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of interests in dinosaurs in the United States, known as dinosaur mania.

Dinosaur mania was exemplified by the fierce rivalry between Edward Drinker Cope and Othniel Charles Marsh, both of whom raced to be the first to find new dinosaurs in what came to be known as the Bone Wars. This fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their diggers often used dynamite to unearth bones. Modern paleontologists would find such methods crude and unacceptable, since blasting easily destroys fossil and stratigraphic evidence. Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast: Marsh unearthed 86 new species of dinosaur and Cope discovered 56, a total of 142 new species. Cope's collection is now at the American Museum of Natural History in New York City, while Marsh's is at the Peabody Museum of Natural History at Yale University.

"Dinosaur renaissance" and beyond

Main article: Dinosaur renaissance
John Ostrom's original restoration of Deinonychus, published in 1969

World War II caused a pause in palaeontological research; after the war, research attention was also diverted increasingly to fossil mammals rather than dinosaurs, which were seen as sluggish and cold-blooded. At the end of the 1960s, however, the field of dinosaur research experienced a surge in activity that remains ongoing. Several seminal studies led to this activity. First, John Ostrom discovered the bird-like dromaeosaurid theropod Deinonychus and described it in 1969. Its anatomy indicated that it was an active predator that was likely warm-blooded, in marked contrast to the then-prevailing image of dinosaurs. Concurrently, Robert T. Bakker published a series of studies that likewise argued for active lifestyles in dinosaurs based on anatomical and ecological evidence (see § Physiology), which were subsequently summarized in his 1986 book The Dinosaur Heresies.

Paleontologist Robert T. Bakker with a mounted skeleton of a tyrannosaurid (Gorgosaurus libratus)

New revelations were supported by an increase in dinosaur discoveries. Major new dinosaur discoveries have been made by paleontologists working in previously unexplored regions, including India, South America, Madagascar, Antarctica, and most significantly China. Across theropods, sauropodomorphs, and ornithischians, the number of named genera began to increase exponentially in the 1990s. In 2008 over 30 new species of dinosaurs were named each year. At least sauropodomorphs experienced a further increase in the number of named species in the 2010s, with an average of 9.3 new species having been named each year between 2009 and 2020. As a consequence, more sauropodomorphs were named between 1990 and 2020 than in all previous years combined. These new localities also led to improvements in overall specimen quality, with new species being increasingly named not on scrappy fossils but on more complete skeletons, sometimes from multiple individuals. Better specimens also led to new species being invalidated less frequently. Asian localities have produced the most complete theropod specimens, while North American localities have produced the most complete sauropodomorph specimens.

Prior to the dinosaur renaissance, dinosaurs were mostly classified using the traditional rank-based system of Linnaean taxonomy. The renaissance was also accompanied by the increasingly widespread application of cladistics, a more objective method of classification based on ancestry and shared traits, which has proved tremendously useful in the study of dinosaur systematics and evolution. Cladistic analysis, among other techniques, helps to compensate for an often incomplete and fragmentary fossil record. Reference books summarizing the state of dinosaur research, such as David B. Weishampel and colleagues' The Dinosauria, made knowledge more accessible and spurred further interest in dinosaur research. The release of the first and second editions of The Dinosauria in 1990 and 2004, and of a review paper by Paul Sereno in 1998, were accompanied by increases in the number of published phylogenetic trees for dinosaurs.

Soft tissue and molecular preservation

An Edmontosaurus specimen's skin impressions found in 1999

Dinosaur fossils are not limited to bones, but also include imprints or mineralized remains of skin coverings, organs, and other tissues. Of these, skin coverings based on keratin proteins are most easily preserved because of their cross-linked, hydrophobic molecular structure. Fossils of keratin-based skin coverings or bony skin coverings are known from most major groups of dinosaurs. Dinosaur fossils with scaly skin impressions have been found since the 19th century. Samuel Beckles discovered a sauropod forelimb with preserved skin in 1852 that was incorrectly attributed to a crocodile; it was correctly attributed by Marsh in 1888 and subject to further study by Reginald Hooley in 1917. Among ornithischians, in 1884 Jacob Wortman found skin impressions on the first known specimen of Edmontosaurus annectens, which were largely destroyed during the specimen's excavation. Owen and Hooley subsequently described skin impressions of Hypsilophodon and Iguanodon in 1885 and 1917. Since then, scale impressions have been most frequently found among hadrosaurids, where the impressions are known from nearly the entire body across multiple specimens.

Color restoration of SinosauropteryxColor restoration of Psittacosaurus

Starting from the 1990s, major discoveries of exceptionally preserved fossils in deposits known as conservation Lagerstätten contributed to research on dinosaur soft tissues. Chiefly among these were the rocks that produced the Jehol (Early Cretaceous) and Yanliao (Mid-to-Late Jurassic) biotas of northeastern China, from which hundreds of dinosaur specimens bearing impressions of feather-like structures (both closely related to birds and otherwise, see § Origin of birds) have been described by Xing Xu and colleagues. In living reptiles and mammals, pigment-storing cellular structures known as melanosomes are partially responsible for producing colouration. Both chemical traces of melanin and characteristically shaped melanosomes have been reported from feathers and scales of Jehol and Yanliao dinosaurs, including both theropods and ornithischians. This has enabled multiple full-body reconstructions of dinosaur colouration, such as for Sinosauropteryx and Psittacosaurus by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur fossils from other localities. (However, some researchers have also suggested that fossilized melanosomes represent bacterial remains.) Stomach contents in some Jehol and Yanliao dinosaurs closely related to birds have also provided indirect indications of diet and digestive system anatomy (e.g., crops). More concrete evidence of internal anatomy has been reported in Scipionyx from the Pietraroja Plattenkalk of Italy. It preserves portions of the intestines, colon, liver, muscles, and windpipe.

Scipionyx fossil with intestines, Natural History Museum of Milan

Concurrently, a line of work led by Mary Higby Schweitzer, Jack Horner, and colleagues reported various occurrences of preserved soft tissues and proteins within dinosaur bone fossils. Various mineralized structures that likely represented red blood cells and collagen fibres had been found by Schweitzer and others in tyrannosaurid bones as early as 1991. However, in 2005, Schweitzer and colleagues reported that a femur of Tyrannosaurus preserved soft, flexible tissue within, including blood vessels, bone matrix, and connective tissue (bone fibers) that had retained their microscopic structure. This discovery suggested that original soft tissues could be preserved over geological time, with multiple mechanisms having been proposed. Later, in 2009, Schweitzer and colleagues reported that a Brachylophosaurus femur preserved similar microstructures, and immunohistochemical techniques (based on antibody binding) demonstrated the presence of proteins such as collagen, elastin, and laminin. Both specimens yielded collagen protein sequences that were viable for molecular phylogenetic analyses, which grouped them with birds as would be expected. The extraction of fragmentary DNA has also been reported for both of these fossils, along with a specimen of Hypacrosaurus. In 2015, Sergio Bertazzo and colleagues reported the preservation of collagen fibres and red blood cells in eight Cretaceous dinosaur specimens that did not show any signs of exceptional preservation, indicating that soft tissue may be preserved more commonly than previously thought. Suggestions that these structures represent bacterial biofilms have been rejected, but cross-contamination remains a possibility that is difficult to detect.

Evolutionary history

Origins and early evolution

Full skeleton of an early carnivorous dinosaur, displayed in a glass case in a museum
The early dinosaurs Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull, from the Triassic

Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic epochs, roughly 20 million years after the devastating Permian–Triassic extinction event wiped out an estimated 96% of all marine species and 70% of terrestrial vertebrate species approximately 252 million years ago. The oldest dinosaur fossils known from substantial remains date to the Carnian epoch of the Triassic period and have been found primarily in the Ischigualasto and Santa Maria Formations of Argentina and Brazil, and the Pebbly Arkose Formation of Zimbabwe.

The Ischigualasto Formation (radiometrically dated at 231–230 million years old) has produced the early saurischian Eoraptor, originally considered a member of the Herrerasauridae but now considered to be an early sauropodomorph, along with the herrerasaurids Herrerasaurus and Sanjuansaurus, and the sauropodomorphs Chromogisaurus, Eodromaeus, and Panphagia. Eoraptor's likely resemblance to the common ancestor of all dinosaurs suggests that the first dinosaurs would have been small, bipedal predators. The Santa Maria Formation (radiometrically dated to be older, at 233.23 million years old) has produced the herrerasaurids Gnathovorax and Staurikosaurus, along with the sauropodomorphs Bagualosaurus, Buriolestes, Guaibasaurus, Macrocollum, Nhandumirim, Pampadromaeus, Saturnalia, and Unaysaurus. The Pebbly Arkose Formation, which is of uncertain age but was likely comparable to the other two, has produced the sauropodomorph Mbiresaurus, along with an unnamed herrerasaurid.

Less well-preserved remains of the sauropodomorphs Jaklapallisaurus and Nambalia, along with the early saurischian Alwalkeria, are known from the Upper Maleri and Lower Maleri Formations of India. The Carnian-aged Chañares Formation of Argentina preserves primitive, dinosaur-like ornithodirans such as Lagosuchus and Lagerpeton in Argentina, making it another important site for understanding dinosaur evolution. These ornithodirans support the model of early dinosaurs as small, bipedal predators. Dinosaurs may have appeared as early as the Anisian epoch of the Triassic, approximately 243 million years ago, which is the age of Nyasasaurus from the Manda Formation of Tanzania. However, its known fossils are too fragmentary to identify it as a dinosaur or only a close relative. The referral of the Manda Formation to the Anisian is also uncertain. Regardless, dinosaurs existed alongside non-dinosaurian ornithodirans for a period of time, with estimates ranging from 5–10 million years to 21 million years.

When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchians, such as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the dinosaurs. Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215 million years ago, a variety of basal archosauromorphs, including the protorosaurs, became extinct. This was followed by the Triassic–Jurassic extinction event (about 201 million years ago), that saw the end of most of the other groups of early archosaurs, like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at least in some areas) at least as late as early –mid Norian and late Norian or earliest Rhaetian stages, respectively, and the exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles. The first few lines of early dinosaurs diversified through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups that became extinct. Also notably, there was a heightened rate of extinction during the Carnian pluvial event.

Evolution and paleobiogeography

The supercontinent Pangaea in the early Mesozoic (around 200 million years ago)

Dinosaur evolution after the Triassic followed changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea, and there was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and early sauropodomorph herbivores. Gymnosperm plants (particularly conifers), a potential food source, radiated in the Late Triassic. Early sauropodomorphs did not have sophisticated mechanisms for processing food in the mouth, and so must have employed other means of breaking down food farther along the digestive tract. The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ceratosaurians, megalosauroids, and allosauroids, and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized metriacanthosaurid theropods and unusual, long-necked sauropods like Mamenchisaurus. Ankylosaurians and ornithopods were also becoming more common, but primitive sauropodomorphs had become extinct. Conifers and pteridophytes were the most common plants. Sauropods, like earlier sauropodomorphs, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential cheek-like organs to keep food in the mouth, and jaw motions to grind food. Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and brachiosaurids through Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid and carcharodontosaurid theropods, and rebbachisaurid and titanosaurian sauropods, also found in South America. In Asia, maniraptoran coelurosaurians like dromaeosaurids, troodontids, and oviraptorosaurians became the common theropods, and ankylosaurids and early ceratopsians like Psittacosaurus became important herbivores. Meanwhile, Australia was home to a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians. The stegosaurians appear to have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of grinding with dental batteries, taken to its extreme in hadrosaurids. Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.

There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosaurids and various types of smaller maniraptoran theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosaurians. In the southern continents that had made up the now-splitting supercontinent Gondwana, abelisaurids were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontid iguanodontians, nodosaurid ankylosaurians, and titanosaurian sauropods were prevalent. Flowering plants were greatly radiating, with the first grasses appearing by the end of the Cretaceous. Grinding hadrosaurids and shearing ceratopsians became very diverse across North America and Asia. Theropods were also radiating as herbivores or omnivores, with therizinosaurians and ornithomimosaurians becoming common.

The Cretaceous–Paleogene extinction event, which occurred approximately 66 million years ago at the end of the Cretaceous, caused the extinction of all dinosaur groups except for the neornithine birds. Some other diapsid groups, including crocodilians, dyrosaurs, sebecosuchians, turtles, lizards, snakes, sphenodontians, and choristoderans, also survived the event.

The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks and chickens, and a variety of waterbirds, diversified rapidly at the beginning of the Paleogene period, entering ecological niches left vacant by the extinction of Mesozoic dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even the larger terrestrial theropods (in the form of Gastornis, eogruiids, bathornithids, ratites, geranoidids, mihirungs, and "terror birds"). It is often stated that mammals out-competed the neornithines for dominance of most terrestrial niches but many of these groups co-existed with rich mammalian faunas for most of the Cenozoic Era. Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals, and ratites are still fairly successful as midsized herbivores; eogruiids similarly lasted from the Eocene to Pliocene, becoming extinct only very recently after over 20 million years of co-existence with many mammal groups.

Classification

Main article: Dinosaur classification Saurischian pelvis structure (left side)Tyrannosaurus pelvis (showing saurischian structure – left side)Ornithischian pelvis structure (left side)Edmontosaurus pelvis (showing ornithischian structure – left side)

Dinosaurs belong to a group known as archosaurs, which also includes modern crocodilians. Within the archosaur group, dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodilians sprawl out to either side.

Collectively, dinosaurs as a clade are divided into two primary branches, Saurischia and Ornithischia. Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all taxa sharing a more recent common ancestor with Triceratops than with Saurischia. Anatomically, these two groups can be distinguished most noticeably by their pelvic structure. Early saurischians—"lizard-hipped", from the Greek sauros (σαῦρος) meaning "lizard" and ischion (ἰσχίον) meaning "hip joint"—retained the hip structure of their ancestors, with a pubis bone directed cranially, or forward. This basic form was modified by rotating the pubis backward to varying degrees in several groups (Herrerasaurus, therizinosauroids, dromaeosaurids, and birds). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups).

By contrast, ornithischians—"bird-hipped", from the Greek ornitheios (ὀρνίθειος) meaning "of a bird" and ischion (ἰσχίον) meaning "hip joint"—had a pelvis that superficially resembled a bird's pelvis: the pubic bone was oriented caudally (rear-pointing). Unlike birds, the ornithischian pubis also usually had an additional forward-pointing process. Ornithischia includes a variety of species that were primarily herbivores.

Despite the terms "bird hip" (Ornithischia) and "lizard hip" (Saurischia), birds are not part of Ornithischia. Birds instead belong to Saurischia, the "lizard-hipped" dinosaurs—birds evolved from earlier dinosaurs with "lizard hips".

Taxonomy

The following is a simplified classification of dinosaur groups based on their evolutionary relationships, and those of the main dinosaur groups Theropoda, Sauropodomorpha and Ornithischia, compiled by Justin Tweet. Further details and other hypotheses of classification may be found on individual articles.

  • Dinosauria
Restoration of six ornithopods; far left: Camptosaurus, left: Iguanodon, center background: Shantungosaurus, center foreground: Dryosaurus, right: Corythosaurus, far right (large) Tenontosaurus.
  • Ornithischia ("bird-hipped"; diverse bipedal and quadrupedal herbivores)
  • Thyreophora (armored dinosaurs; bipeds and quadrupeds)
  • Eurypoda (heavy, quadrupedal thyreophorans)
Restoration of four ceratopsids: top left – Triceratops, top right – Styracosaurus, bottom left – Anchiceratops, bottom right – Chasmosaurus.
  • Ceratopsidae (large, elaborately ornamented ceratopsians)
  • Chasmosaurinae (ceratopsids with enlarged brow horns)
  • Centrosaurinae (ceratopsids mostly characterized by frill and nasal ornamentation)
  • Ornithopoda (various sizes; bipeds and quadrupeds; evolved a method of chewing using skull flexibility and numerous teeth)
  • Elasmaria (mostly southern ornithopods with mineralized plates along the ribs; may be thescelosaurids)
  • Dryomorpha (Dryosaurus and more advanced ornithopods)
  • Hadrosauriformes (ancestrally had a thumb spike; large quadrupedal herbivores, with teeth merged into dental batteries)
Restoration of four macronarian sauropods: from left to right Camarasaurus, Brachiosaurus, Giraffatitan, and Euhelopus
  • Sauropodomorpha (herbivores with small heads, long necks, and long tails)
  • Sauropoda (very large and heavy; quadrupedal)
  • Diplodocoidea (skulls and tails elongated; teeth typically narrow and pencil-like)
  • Dicraeosauridae (small, short-necked diplodocoids with enlarged cervical and dorsal vertebrae)
  • Diplodocidae (extremely long-necked)
  • Macronaria (boxy skulls; spoon- or pencil-shaped teeth)
  • Euhelopodidae (stocky, mostly Asian)
  • Diamantinasauria (horse-like skulls; restricted to the Southern Hemisphere; may be titanosaurs)
  • Titanosauria (diverse; stocky, with wide hips; most common in the Late Cretaceous of southern continents)
  • Coelophysoidea (early theropods; includes Coelophysis and close relatives)
  • †"Dilophosaur-grade neotheropods" (larger kink-snouted dinosaurs)
  • Averostra ("bird snouts")
  • Ceratosauria (generally elaborately horned carnivores that existed from the Jurassic to Cretaceous periods, originally included Coelophysoidea)
  • Abelisauridae (large abelisauroids with short arms and oftentimes elaborate facial ornamentation)
  • Noasauridae (diverse, generally light theropods; may include several obscure taxa)
  • Carnosauria (large meat-eating dinosaurs; megalosauroids sometimes included)
  • Coelurosauria (feathered theropods, with a range of body sizes and niches)
  • Megaraptora? (theropods with large hand claws; potentially tyrannosauroids or neovenatorids)
  • †"Nexus of basal coelurosaurs" (used by Tweet to denote well-known taxa with unstable positions at the base of Coelurosauria)
  • Tyrannoraptora ("tyrant thieves")
  • Ornithomimosauria (small-headed, mostly toothless, omnivorous or possible herbivores)
Restoration of six dromaeosaurid theropods: from left to right Microraptor, Velociraptor, Austroraptor, Dromaeosaurus, Utahraptor, and Deinonychus
  • Caenagnathidae (toothless oviraptorosaurs known from North America and Asia)
  • Oviraptoridae (characterized by two bony projections at the back of the mouth; exclusive to Asia)
  • Paraves (avialans and their closest relatives)
  • Microraptoria (characterized by large wings on both the arms and legs; may have been capable of powered flight)
  • Eudromaeosauria (hunters with greatly enlarged sickle claws)
  • Avialae (modern birds and extinct relatives)

Timeline of major groups

Timeline of major dinosaur groups per Holtz (2007).

QuaternaryNeogenePaleogeneCretaceousJurassicTriassicHolocenePleistocenePlioceneMioceneOligoceneEocenePaleoceneLate CretaceousEarly CretaceousLate JurassicMiddle JurassicEarly JurassicLate TriassicMiddle TriassicEarly TriassicOrnithopodaCeratopsiaPachycephalosauriaAnkylosauriaStegosauriaHeterodontosauridaeAvialaeDeinonychosauriaOviraptorosauriaTherizinosauriaAlvarezsauriaOrnithomimosauriaCompsognathidaeTyrannosauroideaMegaraptoraCarnosauriaMegalosauroideaCeratosauriaCoelophysoideaTitanosauriaBrachiosauridaeDiplodocoideaCetiosauridaeTuriasauriaVulcanodontidaeMassospondylidaeRiojasauridaePlateosauridaeGuaibasauridaeHerrerasauridaeQuaternaryNeogenePaleogeneCretaceousJurassicTriassicHolocenePleistocenePlioceneMioceneOligoceneEocenePaleoceneLate CretaceousEarly CretaceousLate JurassicMiddle JurassicEarly JurassicLate TriassicMiddle TriassicEarly Triassic

Paleobiology

Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces, trackways, gastroliths, feathers, impressions of skin, internal organs and other soft tissues. Many fields of study contribute to our understanding of dinosaurs, including physics (especially biomechanics), chemistry, biology, and the Earth sciences (of which paleontology is a sub-discipline). Two topics of particular interest and study have been dinosaur size and behavior.

Size

Main article: Dinosaur size
Scale diagram comparing the average human to the longest known dinosaurs in five major clades:  Sauropoda (Supersaurus vivianae)  Ornithopoda (Shantungosaurus giganteus)  Theropoda (Spinosaurus aegyptiacus)   Thyreophora (Stegosaurus ungulatus)   Marginocephalia (Triceratops prorsus)

Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic, Late Jurassic and Cretaceous. Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the 100-to-1,000 kg (220-to-2,200 lb) category when sorted by estimated weight into categories based on order of magnitude, whereas recent predatory carnivoran mammals peak in the 10-to-100 kg (22-to-220 lb) category. The mode of Mesozoic dinosaur body masses is between 1 and 10 metric tons (1.1 and 11.0 short tons). This contrasts sharply with the average size of Cenozoic mammals, estimated by the National Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb).

The sauropods were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest was an order of magnitude more massive than anything else that has since walked the Earth. Giant prehistoric mammals such as Paraceratherium (the largest land mammal ever) were dwarfed by the giant sauropods, and only modern whales approach or surpass them in size. There are several proposed advantages for the large size of sauropods, including protection from predation, reduction of energy use, and longevity, but it may be that the most important advantage was dietary. Large animals are more efficient at digestion than small animals, because food spends more time in their digestive systems. This also permits them to subsist on food with lower nutritive value than smaller animals. Sauropod remains are mostly found in rock formations interpreted as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse would have been advantageous in such environments.

Largest and smallest

Scientists will probably never be certain of the largest and smallest dinosaurs to have ever existed. This is because only a tiny percentage of animals were ever fossilized and most of these remain buried in the earth. Few non-avian dinosaur specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork.

Comparative size of Argentinosaurus to the average human

The tallest and heaviest dinosaur known from good skeletons is Giraffatitan brancai (previously classified as a species of Brachiosaurus). Its remains were discovered in Tanzania between 1907 and 1912. Bones from several similar-sized individuals were incorporated into the skeleton now mounted and on display at the Museum für Naturkunde in Berlin; this mount is 12 meters (39 ft) tall and 21.8 to 22.5 meters (72 to 74 ft) long, and would have belonged to an animal that weighed between 30000 and 60000 kilograms (70000 and 130000 lb). The longest complete dinosaur is the 27 meters (89 ft) long Diplodocus, which was discovered in Wyoming in the United States and displayed in Pittsburgh's Carnegie Museum of Natural History in 1907. The longest dinosaur known from good fossil material is Patagotitan: the skeleton mount in the American Museum of Natural History in New York is 37 meters (121 ft) long. The Museo Municipal Carmen Funes in Plaza Huincul, Argentina, has an Argentinosaurus reconstructed skeleton mount that is 39.7 meters (130 ft) long.

Maraapunisaurus, one of the largest animals to walk the earth.
Bruhathkayosaurus, potentially the largest terrestrial animal to ever exist.

There were larger dinosaurs, but knowledge of them is based entirely on a small number of fragmentary fossils. Most of the largest herbivorous specimens on record were discovered in the 1970s or later, and include the massive Argentinosaurus, which may have weighed 80000 to 100000 kilograms (88 to 110 short tons) and reached lengths of 30 to 40 meters (98 to 131 ft); some of the longest were the 33.5-meter (110 ft) long Diplodocus hallorum (formerly Seismosaurus), the 33-to-34-meter (108 to 112 ft) long Supersaurus, and 37-meter (121 ft) long Patagotitan; and the tallest, the 18-meter (59 ft) tall Sauroposeidon, which could have reached a sixth-floor window. There were a few dinosaurs that was considered either the heaviest and longest. The most famous one include Amphicoelias fragillimus, known only from a now lost partial vertebral neural arch described in 1878. Extrapolating from the illustration of this bone, the animal may have been 58 meters (190 ft) long and weighed 122400 kg (269800 lb). However, recent research have placed Amphicoelias from the long, gracile diplodocid to the shorter but much stockier rebbachisaurid. Now renamed as Maraapunisaurus, this sauropod now stands as much as 40 meters (130 ft) long and weigh as much as 120000 kg (260000 lb). Another contender of this title includes Bruhathkayosaurus, a controversial taxon that was recently confirmed to exist after archived photos were uncovered. Bruhathkayosaurus was a titanosaur and would have most likely weighed more than even Marrapunisaurus. Recent size estimates in 2023 have placed this sauropod reaching lengths of up to 44 m (144 ft) long and a colossal weight range of around 110000–170000 kg (240000–370000 lb), if these upper estimates up true, Bruhathkayosaurus would have rivaled the blue whale and Perucetus colossus as one of the largest animals to have ever existed.

The largest carnivorous dinosaur was Spinosaurus, reaching a length of 12.6 to 18 meters (41 to 59 ft) and weighing 7 to 20.9 metric tons (7.7 to 23.0 short tons). Other large carnivorous theropods included Giganotosaurus, Carcharodontosaurus, and Tyrannosaurus. Therizinosaurus and Deinocheirus were among the tallest of the theropods. The largest ornithischian dinosaur was probably the hadrosaurid Shantungosaurus giganteus which measured 16.6 meters (54 ft). The largest individuals may have weighed as much as 16 metric tons (18 short tons).

An adult bee hummingbird, the smallest known dinosaur

The smallest dinosaur known is the bee hummingbird, with a length of only 5 centimeters (2.0 in) and mass of around 1.8 g (0.063 oz). The smallest known non-avialan dinosaurs were about the size of pigeons and were those theropods most closely related to birds. For example, Anchiornis huxleyi is currently the smallest non-avialan dinosaur described from an adult specimen, with an estimated weight of 110 g (3.9 oz) and a total skeletal length of 34 centimeters (1.12 ft). The smallest herbivorous non-avialan dinosaurs included Microceratus and Wannanosaurus, at about 60 centimeters (2.0 ft) long each.

Behavior

A nesting ground of the hadrosaur Maiasaura peeblesorum was discovered in 1978

Many modern birds are highly social, often found living in flocks. There is general agreement that some behaviors that are common in birds, as well as in crocodilians (closest living relatives of birds), were also common among extinct dinosaur groups. Interpretations of behavior in fossil species are generally based on the pose of skeletons and their habitat, computer simulations of their biomechanics, and comparisons with modern animals in similar ecological niches.

The first potential evidence for herding or flocking as a widespread behavior common to many dinosaur groups in addition to birds was the 1878 discovery of 31 Iguanodon, ornithischians that were then thought to have perished together in Bernissart, Belgium, after they fell into a deep, flooded sinkhole and drowned. Other mass-death sites have been discovered subsequently. Those, along with multiple trackways, suggest that gregarious behavior was common in many early dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that duck-billed (hadrosaurids) may have moved in great herds, like the American bison or the African springbok. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in Oxfordshire, England, although there is no evidence for specific herd structures. Congregating into herds may have evolved for defense, for migratory purposes, or to provide protection for young. There is evidence that many types of slow-growing dinosaurs, including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. One example is a site in Inner Mongolia that has yielded remains of over 20 Sinornithomimus, from one to seven years old. This assemblage is interpreted as a social group that was trapped in mud. The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as pack hunters working together to bring down large prey. However, this lifestyle is uncommon among modern birds, crocodiles, and other reptiles, and the taphonomic evidence suggesting mammal-like pack hunting in such theropods as Deinonychus and Allosaurus can also be interpreted as the results of fatal disputes between feeding animals, as is seen in many modern diapsid predators.

Restoration of two Centrosaurus apertus engaged in intra-specific combat

The crests and frills of some dinosaurs, like the marginocephalians, theropods and lambeosaurines, may have been too fragile to be used for active defense, and so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and territorialism. Head wounds from bites suggest that theropods, at least, engaged in active aggressive confrontations.

From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the Gobi Desert in 1971. It included a Velociraptor attacking a Protoceratops, providing evidence that dinosaurs did indeed attack each other. Additional evidence for attacking live prey is the partially healed tail of an Edmontosaurus, a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten by a tyrannosaur but survived. Cannibalism amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus.

Comparisons between the scleral rings of dinosaurs and modern birds and reptiles have been used to infer daily activity patterns of dinosaurs. Although it has been suggested that most dinosaurs were active during the day, these comparisons have shown that small predatory dinosaurs such as dromaeosaurids, Juravenator, and Megapnosaurus were likely nocturnal. Large and medium-sized herbivorous and omnivorous dinosaurs such as ceratopsians, sauropodomorphs, hadrosaurids, ornithomimosaurs may have been cathemeral, active during short intervals throughout the day, although the small ornithischian Agilisaurus was inferred to be diurnal.

Based on fossil evidence from dinosaurs such as Oryctodromeus, some ornithischian species seem to have led a partially fossorial (burrowing) lifestyle. Many modern birds are arboreal (tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines. While some early bird-like species may have already been arboreal as well (including dromaeosaurids) such as Microraptor) most non-avialan dinosaurs seem to have relied on land-based locomotion. A good understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior; the science of biomechanics, pioneered by Robert McNeill Alexander, has provided significant insight in this area. For example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast dinosaurs could run, whether diplodocids could create sonic booms via whip-like tail snapping, and whether sauropods could float.

Communication

Modern birds communicate by visual and auditory signals, and the wide diversity of visual display structures among fossil dinosaur groups, such as horns, frills, crests, sails, and feathers, suggests that visual communication has always been important in dinosaur biology. Reconstruction of the plumage color of Anchiornis suggest the importance of color in visual communication in non-avian dinosaurs. Vocalization in non-avian dinosaurs is less certain. In birds, the larynx plays no role in sound production. Instead, birds vocalize with a novel organ, the syrinx, farther down the trachea. The earliest remains of a syrinx were found in a specimen of the duck-like Vegavis iaai dated 69 –66 million years ago, and this organ is unlikely to have existed in non-avian dinosaurs.

Restoration of a striking and unusual visual display in a Lambeosaurus magnicristatus. The crest may also have acted as a resonating chamber for sounds.

On the basis that non-avian dinosaurs did not have syrinxes and that their next close living relatives, crocodilians, use the larynx, Phil Senter, a paleontologist, has suggested that the non-avians could not vocalize, because the common ancestor would have been mute. He states that they mostly on visual displays and possibly non-vocal sounds, such as hissing, jaw-grinding or -clapping, splashing, and wing-beating (possible in winged maniraptoran dinosaurs). Other researchers have countered that vocalizations also exist in turtles, the closest relatives of archosaurs, suggesting that the trait is ancestral to their lineage. In addition, vocal communication in dinosaurs is indicated by the development of advanced hearing in nearly all major groups. Hence the syrinx may have supplemented and then replaced the larynx as a vocal organ, without a "silent period" in bird evolution.

In 2023, a fossilized larynx was described, from a specimen of the ankylosaurid Pinacosaurus. The structure was composed of cricoid and arytenoid cartilages, similar to those of non-avian reptiles; but the mobile cricoid–arytenoid joint and long arytenoid cartilages would have allowed air-flow control similar to that of birds, and thus could have made bird-like vocalizations. In addition, the cartilages were ossified, implying that laryngeal ossification is a feature of some non-avian dinosaurs. A 2016 study concludes that some dinosaurs may have produced closed-mouth vocalizations, such as cooing, hooting, and booming. These occur in both reptiles and birds and involve inflating the esophagus or tracheal pouches. Such vocalizations evolved independently in extant archosaurs numerous times, following increases in body size. The crests of some hadrosaurids and the nasal chambers of ankylosaurids may have been resonators.

Reproductive biology

See also: Dinosaur egg
Three bluish eggs with black speckling sit atop a layer of white mollusk shell pieces, surrounded by sandy ground and small bits of bluish stone
Nest of a plover (Charadrius)

All dinosaurs laid amniotic eggs. Dinosaur eggs were usually laid in a nest. Most species create somewhat elaborate nests which can be cups, domes, plates, beds scrapes, mounds, or burrows. Some species of modern bird have no nests; the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body and feet. Primitive birds and many non-avialan dinosaurs often lay eggs in communal nests, with males primarily incubating the eggs. While modern birds have only one functional oviduct and lay one egg at a time, more primitive birds and dinosaurs had two oviducts, like crocodiles. Some non-avialan dinosaurs, such as Troodon, exhibited iterative laying, where the adult might lay a pair of eggs every one or two days, and then ensured simultaneous hatching by delaying brooding until all eggs were laid.

When laying eggs, females grow a special type of bone between the hard outer bone and the marrow of their limbs. This medullary bone, which is rich in calcium, is used to make eggshells. A discovery of features in a Tyrannosaurus skeleton provided evidence of medullary bone in extinct dinosaurs and, for the first time, allowed paleontologists to establish the sex of a fossil dinosaur specimen. Further research has found medullary bone in the carnosaur Allosaurus and the ornithopod Tenontosaurus. Because the line of dinosaurs that includes Allosaurus and Tyrannosaurus diverged from the line that led to Tenontosaurus very early in the evolution of dinosaurs, this suggests that the production of medullary tissue is a general characteristic of all dinosaurs.

Fossil interpreted as a nesting oviraptorid Citipati at the American Museum of Natural History.

Another widespread trait among modern birds (but see below in regards to fossil groups and extant megapodes) is parental care for young after hatching. Jack Horner's 1978 discovery of a Maiasaura ("good mother lizard") nesting ground in Montana demonstrated that parental care continued long after birth among ornithopods. A specimen of the oviraptorid Citipati osmolskae was discovered in a chicken-like brooding position in 1993, which may indicate that they had begun using an insulating layer of feathers to keep the eggs warm. An embryo of the basal sauropodomorph Massospondylus was found without teeth, indicating that some parental care was required to feed the young dinosaurs. Trackways have also confirmed parental behavior among ornithopods from the Isle of Skye in northwestern Scotland.

However, there is ample evidence of precociality or superprecociality among many dinosaur species, particularly theropods. For instance, non-ornithuromorph birds have been abundantly demonstrated to have had slow growth rates, megapode-like egg burying behavior and the ability to fly soon after birth. Both Tyrannosaurus and Troodon had juveniles with clear superprecociality and likely occupying different ecological niches than the adults. Superprecociality has been inferred for sauropods.

Genital structures are unlikely to fossilize as they lack scales that may allow preservation via pigmentation or residual calcium phosphate salts. In 2021, the best preserved specimen of a dinosaur's cloacal vent exterior was described for Psittacosaurus, demonstrating lateral swellings similar to crocodylian musk glands used in social displays by both sexes and pigmented regions which could also reflect a signalling function. However, this specimen on its own does not offer enough information to determine whether this dinosaur had sexual signalling functions; it only supports the possibility. Cloacal visual signalling can occur in either males or females in living birds, making it unlikely to be useful to determine sex for extinct dinosaurs.

Physiology

Main article: Physiology of dinosaurs

Because both modern crocodilians and birds have four-chambered hearts (albeit modified in crocodilians), it is likely that this is a trait shared by all archosaurs, including all dinosaurs. While all modern birds have high metabolisms and are endothermic ("warm-blooded"), a vigorous debate has been ongoing since the 1960s regarding how far back in the dinosaur lineage this trait extended. Various researchers have supported dinosaurs as being endothermic, ectothermic ("cold-blooded"), or somewhere in between. An emerging consensus among researchers is that, while different lineages of dinosaurs would have had different metabolisms, most of them had higher metabolic rates than other reptiles but lower than living birds and mammals, which is termed mesothermy by some. Evidence from crocodiles and their extinct relatives suggests that such elevated metabolisms could have developed in the earliest archosaurs, which were the common ancestors of dinosaurs and crocodiles.

This 1897 restoration of Brontosaurus as an aquatic, tail-dragging animal, by Charles R. Knight, typified early views on dinosaur lifestyles.

After non-avian dinosaurs were discovered, paleontologists first posited that they were ectothermic. This was used to imply that the ancient dinosaurs were relatively slow, sluggish organisms, even though many modern reptiles are fast and light-footed despite relying on external sources of heat to regulate their body temperature. The idea of dinosaurs as ectothermic remained a prevalent view until Robert T. Bakker, an early proponent of dinosaur endothermy, published an influential paper on the topic in 1968. Bakker specifically used anatomical and ecological evidence to argue that sauropods, which had hitherto been depicted as sprawling aquatic animals with their tails dragging on the ground, were endotherms that lived vigorous, terrestrial lives. In 1972, Bakker expanded on his arguments based on energy requirements and predator-prey ratios. This was one of the seminal results that led to the dinosaur renaissance.

One of the greatest contributions to the modern understanding of dinosaur physiology has been paleohistology, the study of microscopic tissue structure in dinosaurs. From the 1960s forward, Armand de Ricqlès suggested that the presence of fibrolamellar bone—bony tissue with an irregular, fibrous texture and filled with blood vessels—was indicative of consistently fast growth and therefore endothermy. Fibrolamellar bone was common in both dinosaurs and pterosaurs, though not universally present. This has led to a significant body of work in reconstructing growth curves and modeling the evolution of growth rates across various dinosaur lineages, which has suggested overall that dinosaurs grew faster than living reptiles. Other lines of evidence suggesting endothermy include the presence of feathers and other types of body coverings in many lineages (see § Feathers); more consistent ratios of the isotope oxygen-18 in bony tissue compared to ectotherms, particularly as latitude and thus air temperature varied, which suggests stable internal temperatures (although these ratios can be altered during fossilization); and the discovery of polar dinosaurs, which lived in Australia, Antarctica, and Alaska when these places would have had cool, temperate climates.

Comparison between the air sacs of an abelisaur and a bird

In saurischian dinosaurs, higher metabolisms were supported by the evolution of the avian respiratory system, characterized by an extensive system of air sacs that extended the lungs and invaded many of the bones in the skeleton, making them hollow. Such respiratory systems, which may have appeared in the earliest saurischians, would have provided them with more oxygen compared to a mammal of similar size, while also having a larger resting tidal volume and requiring a lower breathing frequency, which would have allowed them to sustain higher activity levels. The rapid airflow would also have been an effective cooling mechanism, which in conjunction with a lower metabolic rate would have prevented large sauropods from overheating. These traits may have enabled sauropods to grow quickly to gigantic sizes. Sauropods may also have benefitted from their size—their small surface area to volume ratio meant that they would have been able to thermoregulate more easily, a phenomenon termed gigantothermy.

Like other reptiles, dinosaurs are primarily uricotelic, that is, their kidneys extract nitrogenous wastes from their bloodstream and excrete it as uric acid instead of urea or ammonia via the ureters into the intestine. This would have helped them to conserve water. In most living species, uric acid is excreted along with feces as a semisolid waste. However, at least some modern birds (such as hummingbirds) can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. This material, as well as the output of the intestines, emerges from the cloaca. In addition, many species regurgitate pellets, and fossil pellets are known as early as the Jurassic from Anchiornis.

The size and shape of the brain can be partly reconstructed based on the surrounding bones. In 1896, Marsh calculated ratios between brain weight and body weight of seven species of dinosaurs, showing that the brain of dinosaurs was proportionally smaller than in today's crocodiles, and that the brain of Stegosaurus was smaller than in any living land vertebrate. This contributed to the widespread public notion of dinosaurs as being sluggish and extraordinarily stupid. Harry Jerison, in 1973, showed that proportionally smaller brains are expected at larger body sizes, and that brain size in dinosaurs was not smaller than expected when compared to living reptiles. Later research showed that relative brain size progressively increased during the evolution of theropods, with the highest intelligence – comparable to that of modern birds – calculated for the troodontid Troodon.

Origin of birds

Main article: Origin of birds

The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas Henry Huxley. After the work of Gerhard Heilmann in the early 20th century, the theory of birds as dinosaur descendants was abandoned in favor of the idea of them being descendants of generalized thecodonts, with the key piece of evidence being the supposed lack of clavicles in dinosaurs. However, as later discoveries showed, clavicles (or a single fused wishbone, which derived from separate clavicles) were not actually absent; they had been found as early as 1924 in Oviraptor, but misidentified as an interclavicle. In the 1970s, Ostrom revived the dinosaur–bird theory, which gained momentum in the coming decades with the advent of cladistic analysis, and a great increase in the discovery of small theropods and early birds. Of particular note have been the fossils of the Jehol Biota, where a variety of theropods and early birds have been found, often with feathers of some type. Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are now generally accepted to have been their closest ancient relatives. They are most closely allied with maniraptoran coelurosaurs. A minority of scientists, most notably Alan Feduccia and Larry Martin, have proposed other evolutionary paths, including revised versions of Heilmann's basal archosaur proposal, or that maniraptoran theropods are the ancestors of birds but themselves are not dinosaurs, only convergent with dinosaurs.

Feathers

Main article: Feathered dinosaurs
Various feathered non-avian dinosaurs, including Archaeopteryx, Anchiornis, Microraptor and Zhenyuanlong

Feathers are one of the most recognizable characteristics of modern birds, and a trait that was also shared by several non-avian dinosaurs. Based on the current distribution of fossil evidence, it appears that feathers were an ancestral dinosaurian trait, though one that may have been selectively lost in some species. Direct fossil evidence of feathers or feather-like structures has been discovered in a diverse array of species in many non-avian dinosaur groups, both among saurischians and ornithischians. Simple, branched, feather-like structures are known from heterodontosaurids, primitive neornithischians, and theropods, and primitive ceratopsians. Evidence for true, vaned feathers similar to the flight feathers of modern birds has been found only in the theropod subgroup Maniraptora, which includes oviraptorosaurs, troodontids, dromaeosaurids, and birds. Feather-like structures known as pycnofibres have also been found in pterosaurs.

However, researchers do not agree regarding whether these structures share a common origin between lineages (i.e., they are homologous), or if they were the result of widespread experimentation with skin coverings among ornithodirans. If the former is the case, filaments may have been common in the ornithodiran lineage and evolved before the appearance of dinosaurs themselves. Research into the genetics of American alligators has revealed that crocodylian scutes do possess feather-keratins during embryonic development, but these keratins are not expressed by the animals before hatching. The description of feathered dinosaurs has not been without controversy in general; perhaps the most vocal critics have been Alan Feduccia and Theagarten Lingham-Soliar, who have proposed that some purported feather-like fossils are the result of the decomposition of collagenous fiber that underlaid the dinosaurs' skin, and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with dinosaurs. However, their views have for the most part not been accepted by other researchers, to the point that the scientific nature of Feduccia's proposals has been questioned.

Archaeopteryx was the first fossil found that revealed a potential connection between dinosaurs and birds. It is considered a transitional fossil, in that it displays features of both groups. Brought to light just two years after Charles Darwin's seminal On the Origin of Species (1859), its discovery spurred the nascent debate between proponents of evolutionary biology and creationism. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, at least one specimen was mistaken for the small theropod Compsognathus. Since the 1990s, a number of additional feathered dinosaurs have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Many of these specimens were unearthed in the lagerstätten of the Jehol Biota. If feather-like structures were indeed widely present among non-avian dinosaurs, the lack of abundant fossil evidence for them may be due to the fact that delicate features like skin and feathers are seldom preserved by fossilization and thus often absent from the fossil record.

Skeleton

Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent another important line of evidence for paleontologists. Areas of the skeleton with important similarities include the neck, pubis, wrist (semi-lunate carpal), arm and pectoral girdle, furcula (wishbone), and breast bone. Comparison of bird and dinosaur skeletons through cladistic analysis strengthens the case for the link.

Soft anatomy

Pneumatopores on the left ilium of Aerosteon riocoloradensis

Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to a 2005 investigation led by Patrick M. O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said. In 2008, scientists described Aerosteon riocoloradensis, the skeleton of which supplies the strongest evidence to date of a dinosaur with a bird-like breathing system. CT scanning of Aerosteon's fossil bones revealed evidence for the existence of air sacs within the animal's body cavity.

Behavioral evidence

Fossils of the troodonts Mei and Sinornithoides demonstrate that some dinosaurs slept with their heads tucked under their arms. This behavior, which may have helped to keep the head warm, is also characteristic of modern birds. Several deinonychosaur and oviraptorosaur specimens have also been found preserved on top of their nests, likely brooding in a bird-like manner. The ratio between egg volume and body mass of adults among these dinosaurs suggest that the eggs were primarily brooded by the male and that the young were highly precocial, similar to many modern ground-dwelling birds.

Some dinosaurs are known to have used gizzard stones like modern birds. These stones are swallowed by animals to aid digestion and break down food and hard fibers once they enter the stomach. When found in association with fossils, gizzard stones are called gastroliths.

Extinction of major groups

Main article: Cretaceous–Paleogene extinction event

All non-avian dinosaurs and most lineages of birds became extinct in a mass extinction event, called the Cretaceous–Paleogene (K-Pg) extinction event, at the end of the Cretaceous period. Above the Cretaceous–Paleogene boundary, which has been dated to 66.038 ± 0.025 million years ago, fossils of non-avian dinosaurs disappear abruptly; the absence of dinosaur fossils was historically used to assign rocks to the ensuing Cenozoic. The nature of the event that caused this mass extinction has been extensively studied since the 1970s, leading to the development of two mechanisms that are thought to have played major roles: an extraterrestrial impact event in the Yucatán Peninsula, along with flood basalt volcanism in India. However, the specific mechanisms of the extinction event and the extent of its effects on dinosaurs are still areas of ongoing research. Alongside dinosaurs, many other groups of animals became extinct: pterosaurs, marine reptiles such as mosasaurs and plesiosaurs, several groups of mammals, ammonites (nautilus-like mollusks), rudists (reef-building bivalves), and various groups of marine plankton. In all, approximately 47% of genera and 76% of species on Earth became extinct during the K-Pg extinction event. The relatively large size of most dinosaurs and the low diversity of small-bodied dinosaur species at the end of the Cretaceous may have contributed to their extinction; the extinction of the bird lineages that did not survive may also have been caused by a dependence on forest habitats or a lack of adaptations to eating seeds for survival.

Pre-extinction diversity

Just before the K-Pg extinction event, the number of non-avian dinosaur species that existed globally has been estimated at between 628 and 1078. It remains uncertain whether the diversity of dinosaurs was in gradual decline before the K-Pg extinction event, or whether dinosaurs were actually thriving prior to the extinction. Rock formations from the Maastrichtian epoch, which directly preceded the extinction, have been found to have lower diversity than the preceding Campanian epoch, which led to the prevailing view of a long-term decline in diversity. However, these comparisons did not account either for varying preservation potential between rock units or for different extents of exploration and excavation. In 1984, Dale Russell carried out an analysis to account for these biases, and found no evidence of a decline; another analysis by David Fastovsky and colleagues in 2004 even showed that dinosaur diversity continually increased until the extinction, but this analysis has been rebutted. Since then, different approaches based on statistics and mathematical models have variously supported either a sudden extinction or a gradual decline. End-Cretaceous trends in diversity may have varied between dinosaur lineages: it has been suggested that sauropods were not in decline, while ornithischians and theropods were in decline.

Impact event

Main article: Chicxulub crater
Luis (left) and his son Walter Alvarez (right) at the K-T Boundary in Gubbio, Italy, 1981
The Chicxulub Crater at the tip of the Yucatán Peninsula; the impactor that formed this crater may have caused the dinosaur extinction.

The bolide impact hypothesis, first brought to wide attention in 1980 by Walter Alvarez, Luis Alvarez, and colleagues, attributes the K-Pg extinction event to a bolide (extraterrestrial projectile) impact. Alvarez and colleagues proposed that a sudden increase in iridium levels, recorded around the world in rock deposits at the Cretaceous–Paleogene boundary, was direct evidence of the impact. Shocked quartz, indicative of a strong shockwave emanating from an impact, was also found worldwide. The actual impact site remained elusive until a crater measuring 180 km (110 mi) wide was discovered in the Yucatán Peninsula of southeastern Mexico, and was publicized in a 1991 paper by Alan Hildebrand and colleagues. Now, the bulk of the evidence suggests that a bolide 5 to 15 kilometers (3 to 9+1⁄2 miles) wide impacted the Yucatán Peninsula 66 million years ago, forming this crater and creating a "kill mechanism" that triggered the extinction event.

Within hours, the Chicxulub impact would have created immediate effects such as earthquakes, tsunamis, and a global firestorm that likely killed unsheltered animals and started wildfires. However, it would also have had longer-term consequences for the environment. Within days, sulfate aerosols released from rocks at the impact site would have contributed to acid rain and ocean acidification. Soot aerosols are thought to have spread around the world over the ensuing months and years; they would have cooled the surface of the Earth by reflecting thermal radiation, and greatly slowed photosynthesis by blocking out sunlight, thus creating an impact winter. (This role was ascribed to sulfate aerosols until experiments demonstrated otherwise.) The cessation of photosynthesis would have led to the collapse of food webs depending on leafy plants, which included all dinosaurs save for grain-eating birds.

Deccan Traps

Main article: Deccan Traps

At the time of the K-Pg extinction, the Deccan Traps flood basalts of India were actively erupting. The eruptions can be separated into three phases around the K-Pg boundary, two prior to the boundary and one after. The second phase, which occurred very close to the boundary, would have extruded 70 to 80% of the volume of these eruptions in intermittent pulses that occurred around 100,000 years apart. Greenhouse gases such as carbon dioxide and sulfur dioxide would have been released by this volcanic activity, resulting in climate change through temperature perturbations of roughly 3 °C (5.4 °F) but possibly as high as 7 °C (13 °F). Like the Chicxulub impact, the eruptions may also have released sulfate aerosols, which would have caused acid rain and global cooling. However, due to large error margins in the dating of the eruptions, the role of the Deccan Traps in the K-Pg extinction remains unclear.

Before 2000, arguments that the Deccan Traps eruptions—as opposed to the Chicxulub impact—caused the extinction were usually linked to the view that the extinction was gradual. Prior to the discovery of the Chicxulub crater, the Deccan Traps were used to explain the global iridium layer; even after the crater's discovery, the impact was still thought to only have had a regional, not global, effect on the extinction event. In response, Luis Alvarez rejected volcanic activity as an explanation for the iridium layer and the extinction as a whole. Since then, however, most researchers have adopted a more moderate position, which identifies the Chicxulub impact as the primary progenitor of the extinction while also recognizing that the Deccan Traps may also have played a role. Walter Alvarez himself has acknowledged that the Deccan Traps and other ecological factors may have contributed to the extinctions in addition to the Chicxulub impact. Some estimates have placed the start of the second phase in the Deccan Traps eruptions within 50,000 years after the Chicxulub impact. Combined with mathematical modelling of the seismic waves that would have been generated by the impact, this has led to the suggestion that the Chicxulub impact may have triggered these eruptions by increasing the permeability of the mantle plume underlying the Deccan Traps.

Whether the Deccan Traps were a major cause of the extinction, on par with the Chicxulub impact, remains uncertain. Proponents consider the climatic impact of the sulfur dioxide released to have been on par with the Chicxulub impact, and also note the role of flood basalt volcanism in other mass extinctions like the Permian-Triassic extinction event. They consider the Chicxulub impact to have worsened the ongoing climate change caused by the eruptions. Meanwhile, detractors point out the sudden nature of the extinction and that other pulses in Deccan Traps activity of comparable magnitude did not appear to have caused extinctions. They also contend that the causes of different mass extinctions should be assessed separately. In 2020, Alfio Chiarenza and colleagues suggested that the Deccan Traps may even have had the opposite effect: they suggested that the long-term warming caused by its carbon dioxide emissions may have dampened the impact winter from the Chicxulub impact.

Possible Paleocene survivors

Non-avian dinosaur remains have occasionally been found above the K-Pg boundary. In 2000, Spencer Lucas and colleagues reported the discovery of a single hadrosaur right femur in the San Juan Basin of New Mexico, and described it as evidence of Paleocene dinosaurs. The rock unit in which the bone was discovered has been dated to the early Paleocene epoch, approximately 64.8 million years ago. If the bone was not re-deposited by weathering action, it would provide evidence that some dinosaur populations survived at least half a million years into the Cenozoic. Other evidence includes the presence of dinosaur remains in the Hell Creek Formation up to 1.3 m (4.3 ft) above the Cretaceous–Paleogene boundary, representing 40,000 years of elapsed time. This has been used to support the view that the K-Pg extinction was gradual. However, these supposed Paleocene dinosaurs are considered by many other researchers to be reworked, that is, washed out of their original locations and then reburied in younger sediments. The age estimates have also been considered unreliable.

Cultural depictions

Main article: Cultural depictions of dinosaurs
Outdated Iguanodon statues created by Benjamin Waterhouse Hawkins for the Crystal Palace Park in 1853
Gertie the Dinosaur (1914) by Winsor McCay, featuring the first animated dinosaur

By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured the popular imagination and become an enduring part of human culture. The entry of the word "dinosaur" into the common vernacular reflects the animals' cultural importance: in English, "dinosaur" is commonly used to describe anything that is impractically large, obsolete, or bound for extinction.

Public enthusiasm for dinosaurs first developed in Victorian England, where in 1854, three decades after the first scientific descriptions of dinosaur remains, a menagerie of lifelike dinosaur sculptures was unveiled in London's Crystal Palace Park. The Crystal Palace dinosaurs proved so popular that a strong market in smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened at parks and museums around the world, ensuring that successive generations would be introduced to the animals in an immersive and exciting way. The enduring popularity of dinosaurs, in its turn, has resulted in significant public funding for dinosaur science, and has frequently spurred new discoveries. In the United States, for example, the competition between museums for public attention led directly to the Bone Wars of the 1880s and 1890s, during which a pair of feuding paleontologists made enormous scientific contributions.

The popular preoccupation with dinosaurs has ensured their appearance in literature, film, and other media. Beginning in 1852 with a passing mention in Charles Dickens' Bleak House, dinosaurs have been featured in large numbers of fictional works. Jules Verne's 1864 novel Journey to the Center of the Earth, Sir Arthur Conan Doyle's 1912 book The Lost World, the 1914 animated film Gertie the Dinosaur (featuring the first animated dinosaur), the iconic 1933 film King Kong, the 1954 Godzilla and its many sequels, the best-selling 1990 novel Jurassic Park by Michael Crichton and its 1993 film adaptation are just a few notable examples of dinosaur appearances in fiction. Authors of general-interest non-fiction works about dinosaurs, including some prominent paleontologists, have often sought to use the animals as a way to educate readers about science in general. Dinosaurs are ubiquitous in advertising; numerous companies have referenced dinosaurs in printed or televised advertisements, either in order to sell their own products or in order to characterize their rivals as slow-moving, dim-witted, or obsolete.

See also

Notes

  1. Dinosaurs (including birds) are members of the natural group Reptilia. Their biology does not precisely correspond to the antiquated class Reptilia of Linnaean taxonomy, consisting of cold-blooded amniotes without fur or feathers. As Linnean taxonomy was formulated for modern animals prior to the study of evolution and paleontology, it fails to account for extinct animals with intermediate traits between traditional classes.

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