2673:(the most common remains found in Dinosaur Park) and noticed that the mandibular symphysis becomes fully ossified relatively early in caeagnathid growth, and it is thus a poor indicator of the precise age of a specimen. Another impediment to precise age identification are the possible irregularities in the deposition of bone tissues as the animals grow. Funston and colleagues notice that the lines of arrested growth (or LAGs), which usually correspond to slower growth during the winter months, may not have been formed in caenagnathids under 1-2 years of age, which may lead to the underestimation of the ages of some specimens.
3077:
903:
1278:
2923:
2584:
1990:
1446:
5281:
2650:
5286:
131:
1063:
2795:
2135:
1591:
2044:
1500:
2017:
1473:
5309:
2074:
1530:
860:, Nicholas Longrich, Ken Barnes, Scott Clark, and Larry Millar, referred ROM 781 to their newly described genus. Their reasoning for this reassignment was the referral of additional specimens — TMP 1992.36.390, TMP 1979.8.622, TMP 1991.144.1 (all lower jaw fragments), and TMP 1982.39.4 (a partial tarsometatarsus) — to the species based on their small size in comparison with all other described caenagnathids.
720:
2344:
1175:
2253:
709:
553:
109:
2198:
1730:
2106:
1562:
2430:
2606:, is described as the "small, robust morphotype" and is distinct from all other caenagnathid pelves in the formation. The most immediately visible distinction is that the pelves of the small, robust morph are curved into a slight letter-s shape when viewed from above or below. Other differences are present but less obvious such as the very small
2306:
1637:
2626:. The similar ontogenetic age of differently-sized postcranial remains indicated the probable presence of three caenagnathid taxa in Dinosaur Park. Two of these taxa had already been named, and the remaining group of specimens — comprising the small, robust morph — were morphologically distinct from the Texan species of
2759:
due to the coossification of its metatarsals and expanded attachment sites for leg muscles along the iliac crest. It is not certain what the ecological implications of these adaptations may have been, but they appear to be unique in that regard among the caenagnathids of the
Dinosaur Park Formation.
1418:
For specimens which do not contain the relevant diagnostic characters or any indication of their ontogenetic age, they have been assigned to existing taxa according to size and locality. Longrich and colleagues note that this approach has shortcomings, but they argue that the ecological abundance of
2734:
While caenagnathid remains are relatively uncommon, they are widely dispersed across
Dinosaur Provincial Park, with no clear pattern of which genera are found at specific localities. This suggests that they were present throughout the faunal zones of the Dinosaur Park Formation, which implies there
2897:
In their description of TMP 2009.12.14, Therrien and colleagues conducted a regression analysis of the relationship between the masses of extant reptilian and mammalian predators and the masses of potential sympatric prey items. They used this, along with the estimated masses of juvenile and adult
1223:
is also preserved on the lateral surface of the centrum. Despite the fact that these bones were not subject to a histological analysis, Funston believes they belong to an adult due to the large muscle attachment sites visible on the bone and the robustness of the iliac crest. The small size in an
1030:
was relatively small in life, with adults measuring 1–2 metres (3.3–6.6 ft) long. Rubén Molina-Pérez and Asier
Larramendi suggested a more specific estimate of 2.1 metres (6.9 ft) long and 82 centimetres (2.69 ft) tall at the hip with an estimated mass of 32 kilograms (71 lb).
2730:
can be divided into several discrete faunal assemblages which were not all contemporaneous. However, it is presently unclear if any of the known caenagnathid genera are restricted to one or more of these faunal zones. Caenagnathid remains have not been recovered as often as those of other
1395:, which allows researchers to diagnose genera and species based on very incomplete remains, but which also means that more complete remains are undiagnostic if they do not preserve the mandible or manus. However, the authors note that the shape of the beak in modern
4041:
Funston, Gregory F.; Wilkinson, Ryan D.; Simon, D. Jade; Leblanc, Aaron H.; Wosik, Mateusz; Currie, Philip J. (2020). "Histology of
Caenagnathid (Theropoda, Oviraptorosauria) Dentaries and Implications for Development, Ontogenetic Edentulism, and Taxonomy".
1411:, so they argue that it is not unreasonable to assume that this may be sufficient in naming new genera or species. Furthermore, the degree of fusion exhibited by the jaw bones, as well as their surface texture, can be indicative of ontogenetic age in many
1042:
including: coossification of the distal ends of tarsals III and IV, a lack of poterior protrusions on the proximal ends of metatarsals II and IV, and a prominent ridge on the posterior surface of metatarsal III. These autapomorphies not only distinguish
2956:
is composed of sediments that were derived from the erosion of the mountains to the west. It was deposited on an alluvial to coastal plain by river systems that flowed eastward and southeastward to the
Bearpaw Sea, a large inland sea that was part of the
986:
was assigned to one of these three genera based on size and diagnoses were amended from the apomorphies of those specimens. Some specimens were not able to be referred to any of these genera with certainty due to the incompleteness of the remains.
1419:
adult animals in any ecosystem means that most fossilized animals will be adults anyways, which they suggest is sufficient for the purposes of their analysis. A consensus of the 116 most parsimonious trees in their analysis is shown below.
2786:, on the other hand, has a sharply upturned tip of the lower jaw, which is possibly an adaptation for a more carnivorous mode of life, although this remains speculative and untestable due to the incompleteness of these remains.
1093:. It consists of three metatarsals which are bowed slightly inwards such that metatarsals II and IV meet at their proximal ends. The length of the metatarsals is stated by Parks as roughly 160 millimetres (6.3 in).
1199:
based solely on the size and locality of the specimen (in comparison to ROM 781) was unfounded, and he states that there is insufficient evidence to suggest that TMP 1992.36.290 belongs to the same taxon as ROM 781.
2660:
The ontogeny of caenagnathids is difficult to study due to the incompleteness of known remains. This is further complicated by the relative lack of overlapping skeletal material from the various named taxa of the
1171:— exemplified by the specimens TMP 1992.36.390, TMP 1979.8.622, and TMP 1991.144.1 — differ substantially from other caenagnathids in their relatively short length and the upturned nature of the dentary symphysis.
1303:. Their analysis used the dataset of Longrich's earlier publication in 2010 with several new taxa added for a total of 28 taxa coded for 205 characters. Their data set also included recent information about
3301:
The
Dinosaur Park Formation also preserves a wide array of smaller animals which would have been abundant in the waterways of the depositional environment. These included a variety turtles such as the large
1105:
as ROM 781 and the diagnosis of the holotype was amended. A re-examination of the holotype led to the discovery that the metatarsals II and IV were coossified and that their separation in the holotype was
2617:
Rhodes and colleagues suggested that these morphological distinctions in the hip bones of caenagnathids were unlikely to be the result of ontogeny because the level of coossification between the ilia and
3748:
Longrich, Nicholas R.; Barnes, Ken; Clark, Scott; Millar, Larry (2013). "Correction to "Caenagnathidae from the Upper
Campanian Aguja Formation of West Texas, and a Revision of the Caenagnathinae"".
1260:
were described by
Funston, but they were not referred to any particular genus within caenagnathidae. These remains, which included jaw bones, were small in size, but similar in morphology to
956:
each of the specimens were in and whether or not the smaller caenagnathid remains belonged to juveniles or adults of smaller species. Funston's analysis resulted in revised diagnoses for
1034:
In his comprehensive re-description of the caenagnathids from the
Dinosaur Park Formation, Gregory Funston provides amended diagnoses for all three named genera in his analysis. For
1219:. However, the anterior articular surface of the centrum is unbroken, which Funston suggests means that the sacrum was not fully fused when the animal died. Evidence of the sacral
3418:
2949:
is not known confidently known. However, the
Dinosaur Park Formation as a whole is relatively confidently known to have been deposited between 77.03 and 75.46 million years ago.
2859:
bones showed a complete lack of growth marks, leading Therrien and colleagues to suggest that these individuals were within their first year of life when they were eaten by the
2751:
is the variation in their adult size. This is the primary means to distinguish remains of the three genera, but there are also several more subtle morphological distinctions.
792:
The classification of ROM 781 was amended again in 1997 by Hans-Dieter Sues in 1997 when he published an extensive monograph describing a newly-discovered specimen of the genus
1026:
assigned to the genus indicated that the individuals that the bones came from were close to adulthood, but not yet fully matured. Funston is confident in his assessment that
5435:
1167:) due to its small size, which they believed was not reflective of its ontogenetic age because of the fusion of the tarsal elements. They also remark that the dentaries of
4273:"Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America"
4212:
Therrien, François; Zelenitsky, Darla K.; Tanaka, Kohei; Voris, Jared T.; Erickson, Gregory M.; Currie, Philip J.; Debuhr, Christopher L.; Kobayashi, Yoshitsugu (2023).
2882:
because the two specimens are known to have been consumed at different times due to their relative locations within the presumed digestive tract of the predator and the
1346:, but there was less resolution in the more derived area of the tree. In particular, their strict consensus tree did not support the unambiguous monophyly of the genus
4487:
2851:
was sectioned and subjected to histological analysis, which revealed that it was between 5-7 years-old when it died and was only about one-eighth the mass of an adult
4129:
Currie, Philip J.; Godfrey, Stephen J.; Nessov, Lev (1993). "New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia".
1322:
of caenagnathidae was supported in their analysis based on the following diagnostic characters: fused dentaries, a ventrally bowed process on the underside of the
2636:
as sister taxa, but there have not been any comprehensive reviews of Funston's phylogenetic analysis to date, nor have any new analyses been conducted as of yet.
4164:
Funston, Gregory F.; Currie, Philip J. (2014). "A previously undescribed caenagnathid mandible from the late Campanian of Alberta, and insights into the diet of
3998:
Rhodes, Matthew M.; Funston, Gregory F.; Currie, Philip J. (2020). "New material reveals the pelvic morphology of Caenagnathidae (Theropoda, Oviraptorosauria)".
2602:
in 2019, Matthew Rhodes and colleagues categorize every specimen into one of three categories. The morph which they believe belongs to a new taxon, later named
1195:, however this assignment is somewhat uncertain because of the lack of overlapping material with the type series. Funston also argues that the designation of a
921:
in order to determine their actual diversity in that depositional environment. Funston personally examined and scored material in the collections of the
5440:
4570:
1289:
analysis to include ROM 781 was the one conducted by Nicholas Longrich, Ken Barnes, Scott Clark, and Larry Millar in their paper describing the genus
761:, and a process on the distal ends of metatarsals II and IV. At the time of this publication, there was an ongoing debate about the validity of the
5376:
3357:— being known from several genera each. The freshwater systems of the region would have also supported a great variety of fish species including
3849:
Barsbold, R. (1976). "О новом позднемеловом семействе мелких теропод Oviraptoridae fam. nov Монголии" [A new Late Cretaceous family of small
2890:
or they would not have been preserved. This, taken together, suggests that the predator may have had a preference for eating the hind limbs of
5460:
1963:. However, Fuston is careful to note that the topology of his tree is not strongly supported and is likely to be revised by future analyses.
4420:
4385:
5445:
2713:
in the specimen, which suggests that the animal survived for at least a short time after it received the injury that caused this fracture.
3915:. Translated by Connolly, David; Ramírez Cruz, Gonzalo Ángel. Illustrated by Andrey Atuchin and Sante Mazzei. Princeton University Press.
2701:
peduncles, and has resulted in a significant swelling of the ilium, which can be clearly seen from above and slightly obscures the brevis
1264:. Funston argues that they cannot be referred to a particular genus without being sectioned and having their ontogenetic age determined.
3936:
Funston, Gregory F.; Chinzorig, Tsogtbaatar; Tsogtbaatar, Khishigjav; Kobayashi, Yoshitsugu; Sullivan, Corwin; Currie, Philip J. (2020).
5302:
3433:
2731:
coelurosaurs, but this is partially accounted for by their relatively small size and lack of teeth, which may present a sampling bias.
5465:
3920:
671:
4438:"A new elasmosaurid (Sauropterygia: Plesiosauria) from the non-marine to paralic Dinosaur Park Formation of southern Alberta, Canada"
4214:"Exceptionally preserved stomach contents of a young tyrannosaurid reveal an ontogenetic dietary shift in an iconic extinct predator"
4353:
1959:
were also recovered as caenagnathids in his analysis, suggesting that the group persisted in Asia even after the diversification of
641:, were believed to be ornithomimids. It would not be until 1976 that oviraptorosaurs became recognized as their own unique clade.
2972:
The Dinosaur Park Formation is roughly 75 metres (250 ft) thick. The lower portion of the formation was laid down in fluvial
5455:
3413:
1415:, which can be used to determine if an individual specimen is a new taxon or simply a different life stage of an existing taxon.
840:
by the presence of co-ossification of the second and fourth metatarsals, and it was included in the genus because Sues regarded
3498:"Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: anatomy, osteohistology, taxonomy, and evolution"
3423:
2735:
was some sort of niche differentiation between the three taxa. The most immediately obvious morphological difference between
1101:
In 1989, the specimens ROM 37163 (distal metatarsal II) and TMP 82.39.4 (a partial tarsometatarsus) were assigned to the same
5280:
4563:
3391:
3386:
2811:
1380:
because of the similar size of the two species, but noted that they may belong to separate genera, pending further research.
536:
1928:
to be more widely distributed throughout the trees in his analysis, rather than grouping together as a single radiation of
2902:
to predict the possible range of prey sizes for young and adult individuals of that genus. Their findings indicated that
938:
684:. He also referred two new specimens to the species. The first of these new specimens, ROM 37163, consists of a partial
2828:
individuals were flexed and contorted into a very compact shape, which suggests that they were inside the stomach of the
1006:
caenagnathids, Gregory Funston concluded that it had a substantially smaller adult size than the contemporaneous genera
5285:
945:
samples from several bones in order to examine them for "lines of arrested growth", which can be used to estimate the
1253:
were noted among these bones, which was reflected in the phylogenetic analysis that was conducted in the same paper.
1938:, which is one of the oldest unambiguous caenagnathids, in a relatively derived position as the sister genus of the
3076:
3542:
Parks, William A. (1933). "New Species of Dinosaurs and Turtles from the Upper Cretaceous Formations of Alberta".
2622:
is similar in very differently-sized specimens. Funston elaborated on this finding in 2020 in his description of
902:
5367:
4556:
922:
2926:
A life reconstruction of several dinosaurs in the fluvial environment that the Dinosaur Park Formation contained
968:, which Funston argues are distinct and valid genera. This analysis also resulted in the specimens assigned to
130:
5312:
4667:
2934:
2693:
that was fractured at some point during the animal's life. The fracture extends along the surface between the
907:
372:
3204:
directly in addition to larger dinosaurs. These large tyrannosaurids were accompanied by an array of smaller
2814:, François Therrien and colleagues described TMP 2009.12.14 — an exceptionally preserved juvenile specimen of
1295:
in 2013. In their analysis, they assign ROM 781 and several other specimens to a second species in the genus
4078:
3706:"Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a Revision of the Caenagnathinae"
3224:. These smaller-bodied theropods are known from substantially less complete remains with some taxa, such as
5450:
4344:
4095:
3185:
3093:
3033:
3030:
2958:
2953:
2946:
2727:
2662:
2599:
1885:
1003:
983:
918:
697:
584:. However, it would be another seven years before a full description of this material was published in the
581:
456:
1277:
5430:
5329:
3400:
3396:
3193:
3189:
758:
2870:
individuals do not appear to have been eaten in their entirety. Only the hind limbs and elements of the
2776:, and it is believed to be primarily herbivorous because the better-understood and closely-related taxon
5402:
4952:
4481:
2628:
1902:, and it included all described oviraptorosaur taxa known from that time as well as a few unnamed taxa.
1683:
1291:
1055:
970:
934:
856:
622:
585:
569:
531:
4096:"Megaherbivorous dinosaur turnover in the Dinosaur Park Formation (upper Campanian) of Alberta, Canada"
696:, and it was collected in 1982 by Linda Strong-Watson. Both of these specimens were discovered in the
2922:
4721:
4518:
4284:
4225:
4177:
4138:
4103:
4007:
3949:
3885:
3659:
3617:
3573:
2595:
1775:
1281:
A phylogenetic analysis showing changes in body size in caenagnathids (top) and oviraptorids (bottom)
1111:
926:
689:
593:
276:
251:
4083:(PhD in Systematics and Evolution thesis). Department of Biological Sciences, University of Alberta.
2583:
894:, by the presence of a strongly upturned beak tip and a chin that is square-shaped in lateral view.
4843:
2973:
2832:, rather than being preserved alongside it or washed together post-mortem. This demonstrated that
2632:
which they had been named as in 2013. Phylogenetic analyses have never found the two "species" of
1327:
1257:
930:
4694:
4534:
4023:
3765:
3725:
3303:
1191:
During his comprehensive analysis in 2020, Gregory Funston referred two partial hip specimens to
309:
125:
5407:
4412:
4406:
1896:. This analysis used the same data set which he developed for the description of the new genus
4377:
4371:
2649:
1989:
1445:
1239:
The remaining specimens, UALVP 59606, UALVP 55585, and UALVP 59606, consist of various partial
5389:
4871:
4836:
4469:
4416:
4402:
4381:
4349:
4312:
4253:
4059:
3975:
3916:
3798:
3519:
3244:
2966:
1212:
1062:
577:
5394:
4367:
1243:
of varying completeness and states of coossification. The latter of these was sectioned for
952:
There has historically been significant uncertainty in the scientific literature as to which
5056:
4789:
4767:
4733:
4579:
4526:
4459:
4449:
4302:
4292:
4243:
4233:
4185:
4146:
4111:
4051:
4015:
3965:
3957:
3893:
3829:
3788:
3757:
3717:
3667:
3625:
3581:
3509:
3354:
2794:
2391:
1970:
1943:
1893:
1426:
1135:
In 2013, Nicholas Longrich and colleagues referred additional material to the newly renamed
1125:
975:, as well as some recently discovered specimens, being referred to the newly-erected genus,
667:
626:
482:
406:
364:
2134:
1590:
4981:
4894:
4878:
4652:
3342:
3217:
3209:
3170:
3106:
2871:
2710:
2702:
2587:
Known caenagnathid remains from the Dinosaur Park Formation shown to scale with each other
2056:
2043:
1512:
1499:
1392:
1331:
1240:
1140:
1121:
1066:
Comparison of the metatarsal elements of several caenagnathids, including the holotype of
693:
618:
453:
30:
3564:
Russell, Dale A. (1972). "Ostrich Dinosaurs from the Late Cretaceous of Western Canada".
4522:
4288:
4248:
4229:
4213:
4181:
4142:
4107:
4011:
3953:
3889:
3663:
3650:, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America".
3621:
3577:
1139:. These specimens included several bones of the lower jaws as well as newly discovered
5249:
5235:
5042:
5019:
4904:
4829:
4814:
4502:
4464:
4437:
4307:
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2086:
2016:
1984:
1934:
1614:
1542:
1472:
1440:
1412:
1216:
817:
654:
588:'s geology journal in 1933 by Dr. William A. Parks. The material, consisting of three
573:
513:
331:
214:
4436:
Campbell, Mitchel, Ryan and Anderson, James A., Mark T., Michael J., Jason S. (2021).
3876:(Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada".
2073:
1529:
688:
and was collected sometime between the years of 1920 and 1954 by an expedition of the
5424:
5221:
5199:
5140:
5124:
5071:
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4931:
4855:
4538:
4027:
3769:
3729:
3705:
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3221:
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2012:
1960:
1918:
1724:
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1525:
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1407:
1368:
1336:
1286:
1014:
964:
767:
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637:
525:
494:
357:
66:
2910:
ranges, juvenile and adult, were within the size range that both juvenile and adult
2343:
5192:
5148:
5106:
5049:
4992:
4966:
4636:
3783:
Osborn, H. F. (1924). "Three new Theropoda, Protoceratops zone, central Mongolia".
3671:
3358:
3334:
3281:
3262:
3257:
3226:
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3134:
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1208:
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914:
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748:
719:
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468:
446:
319:
259:
227:
4530:
4019:
3088:(left) in their natural environment with other dinosaurs visible in the background
2961:. That sea gradually inundated the adjacent coastal plain, depositing the marine
2252:
5352:
4297:
4115:
1334:. The analysis did resolve several taxa at the base of caenagnathidae including
674:. Currie referred the specimen to a new species in the recently-described genus
5361:
5242:
5175:
5113:
5035:
5028:
4864:
4706:
3271:
3174:
2816:
2778:
2424:
2337:
1924:
1401:
1372:, and the specimen BMNH 2033 (which was referred as an indeterminate species of
1305:
349:
41:
4080:
Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda)
2933:
was discovered at the Little Sand Hill Creek locality, which is located within
2197:
1729:
1155:
of the species. In their re-assignment, Longrich and colleagues distinguished
708:
552:
108:
5256:
5207:
5084:
5008:
4938:
4924:
4917:
4618:
4609:
3938:"A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria"
3374:
3350:
3290:
3286:
3252:
3148:
3065:
3053:
3018:
2988:
2894:, which are hypothesized to have been more muscular and thus more nutritious.
2300:
2158:
2038:
1955:
1905:
This analysis recovered similar results to most prior analyses. Namely, that
1631:
1249:
1220:
1215:
is characteristically low and wide, which is the condition seen in many other
1116:
1049:
1023:
785:
779:
753:
685:
676:
629:
was not yet recognized as a unique clade, and several oviraptorans, including
614:
589:
519:
342:
188:
86:
51:
4342:
Eberth, D.A. 2005. The geology. In: Currie, P.J., and Koppelhus, E.B. (eds),
3523:
5214:
5163:
5156:
5091:
4959:
4821:
4679:
4627:
3802:
3427:
3326:
3307:
3295:
3205:
3061:
3007:
3000:
2996:
2980:
2879:
2706:
2666:
2214:
1939:
1929:
1889:
1700:
1356:
1319:
1244:
1107:
1019:
942:
862:
848:
845:
631:
201:
142:
91:
4473:
4316:
4257:
4238:
4189:
4063:
3979:
2105:
1561:
5346:
5133:
5098:
4973:
4807:
4603:
3850:
3761:
3721:
3370:
3362:
3346:
3341:
were also abundant in the area with representatives from all three major
3318:
3251:
The waterways of the Dinosaur Park Formation also played host to several
3230:, being known only from teeth. More well-known small theropods included
3097:
2992:
2984:
2907:
2670:
2247:
1949:
1494:
1388:
1351:
1323:
1311:
1196:
1152:
1080:
953:
946:
502:
449:
392:
175:
162:
81:
76:
61:
56:
46:
4454:
3961:
3116:, are known from various layers in addition to eight or more species of
2429:
5381:
3514:
3497:
3408:
3404:
3338:
3330:
3045:
3037:
2991:
environments, consists primarily of massive to laminated, organic-rich
2938:
2698:
2305:
1636:
1148:
1120:
included the shape of the bones' cross-sections and the presence of an
486:
460:
439:
96:
71:
3818:"Two new theropod dinosaurs from the Belly River Formation of Alberta"
4597:
4055:
3834:
3817:
3793:
3313:
3261:, has been named. Among the other aquatic reptiles present were the
3013:
These geological features are interpreted as a low-relief setting of
2942:
2619:
2592:
1898:
1224:
adult animal precludes the assignment of TMP 1981.023.0039 to either
1204:
464:
152:
5323:
4507:, gen. et sp. nov., a Late Cretaceous Canadian azhdarchid pterosaur"
4150:
3897:
3872:
Currie, P.J.; Russell, D.A. (1988). "Osteology and relationships of
3629:
3585:
1916:; Funston found several traditionally "elmisaurine" genera, such as
1399:
is an important diagnostic trait, as well as in some dinosaurs like
4548:
4348:. Indiana University Press: Bloomington and Indianapolis, p.54-82.
3913:
Dinosaur Facts and Figures: The Theropods and Other Dinosauriformes
666:
A more thorough re-examination of ROM 781 was conducted in 1988 by
5228:
5184:
4345:
Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed
3075:
3041:
3026:
3022:
3014:
2962:
2921:
2844:
2793:
2648:
2582:
1913:
1276:
1203:
The first of the new specimens, TMP 1981.023.0039, is an anterior
1173:
1102:
1061:
1047:
from its contemporaries, but also from the closely related genera
901:
551:
442:
3100:
assemblages known in the fossil record. At least six species of
1256:
Several other caenagnathid specimens from the collections of the
556:
An outcrop of the Dinosaur Park Formation, where the holotype of
3057:
3003:
2999:. The Lethbridge Coal Zone, which consists of several seams of
2976:
environments and consists primarily of fine- to medium-grained,
1396:
5327:
4590:
4552:
3366:
1247:
analysis. Similarities with the known metatarsal elements of
783:, and Currie used his re-description of ROM 781 to argue that
3704:
Longrich, N. R.; Barnes, K.; Clark, S.; Millar, L. (2013).
2653:
Histological section of one of the metatarsals referred to
2610:
and the expanded dorsal ridge on the posterior part of the
427:
421:
4077:
Funston, Gregory F. (2019). "Chapter 3 — Caenagnathidae".
4405:. In Currie, Philip J.; Koppelhus, Eva Bundgaard (eds.).
4370:. In Currie, Philip J.; Koppelhus, Eva Bundgaard (eds.).
653:
by Dale Russell in 1972. However, this genus was made a
415:
3142:
are also abundant, being represented by members of both
2945:. The age of this locality relative to the rest of the
692:. The second, TMP 82.39.4, consists of another partial
670:, who published a re-description of the material in the
505:
history, and has been previously assigned to the genera
1350:; the two species they assigned to the genus were in a
1315:
which had been published since the analysis in 2010.
1087:
was described in 1933 by William Parks as a species of
700:
and resembled the holotype enough to warrant referral.
3419:
List of stratigraphic units with dinosaur body fossils
3294:
has also been preserved alongside some indeterminate
1383:
Longrich and colleagues broadly determined that most
1376:). The authors remark that they referred ROM 781 to
1159:
from the genera to which it was assigned previously (
854:
In 2013, accompanying their description of the genus
430:
412:
4501:
Hone, D.; Habib, M.; Therrien, F. (September 2019).
424:
418:
292:
266:
5336:
5174:
5123:
5070:
5018:
4991:
4903:
4892:
4854:
4787:
4732:
4705:
4678:
4651:
2689:) is TMP 1992.036.0476, which is a mostly complete
913:In 2020, Gregory Funston conducted a review of all
409:
1326:, the lack of contact between the dentary and the
603:It was named as a new species of the common genus
120:with the material described in 2020 shown in pink
4100:Palaeogeography, Palaeoclimatology, Palaeoecology
3750:Bulletin of the Peabody Museum of Natural History
3710:Bulletin of the Peabody Museum of Natural History
3604:Currie, Philip J. (1989). "The first records of
3544:University of Toronto Studies, Geological Series
3424:Cretaceous land vertebrate ages in North America
3911:Molina-Pérez, Rubén; Larramendi, Asier (2019).
2874:are preserved within the stomach cavity of the
1387:genera are diagnosed by characteristics of the
1182:, which was sectioned for histological analysis
3491:
3489:
3487:
3485:
3483:
3481:
3479:
3477:
3475:
3473:
3471:
3469:
3321:at Dinosaur Park, with taxa from the families
2665:. Gregory Funston and colleagues conducted a
4564:
4401:Braman, Dennis R.; Koppelhus, Eva B. (2005).
3608:(Saurischia, Theropoda) from North America".
3467:
3465:
3463:
3461:
3459:
3457:
3455:
3453:
3451:
3449:
3040:was warmer than present-day Alberta, without
572:'s expeditions to the Sand Creek area of the
564:The material which would eventually be named
8:
4486:: CS1 maint: multiple names: authors list (
3743:
3741:
3739:
3096:preserves one of the most diverse non-avian
2995:with abundant root traces, and thin beds of
2983:. The upper portion, which was deposited in
2886:must have died prior to fully digesting the
2820:, which contained remains from at least two
649:ROM 781 was reassigned to the dubious genus
467:. The genus contains only one species, the
3502:Vertebrate Anatomy Morphology Palaeontology
982:Most of the caenagnathid material from the
757:, by the presence of a weaker ridge on the
5436:Late Cretaceous dinosaurs of North America
5324:
5076:
4909:
4900:
4799:
4795:
4784:
4711:
4684:
4657:
4648:
4587:
4571:
4557:
4549:
3537:
3535:
3533:
3403:- roughly coeval geological formations in
2755:is believed to have been more adapted for
874:were all much larger than the newly named
725:Known hand and foot remains of the genera
107:
20:
4463:
4453:
4306:
4296:
4247:
4237:
3993:
3991:
3989:
3969:
3833:
3792:
3513:
2709:breakage of the fossil has also revealed
1211:(TMP 1981.023.0034-5). The shape of the
886:. Longrich and colleagues distinguished
613:. This name was assigned based on three
568:was discovered in 1926 during one of the
4338:
4336:
4334:
4332:
4330:
4328:
4326:
3853:Oviraptoridae n. fam. in Mongolia].
3641:
3639:
1207:which was found in association with two
844:as the only valid caenagnathid genus in
4207:
4205:
4203:
4201:
4199:
3699:
3697:
3695:
3693:
3691:
3689:
3687:
3685:
3683:
3681:
3559:
3557:
3445:
2969:on top of the Dinosaur Park Formation.
2782:is believed to have been a herbivore.
4479:
3599:
3597:
3595:
3165:Preying on this wide variety of large
2598:of all caenagnathid material from the
617:, which were believed to belong to an
4411:. Indiana University Press. pp.
4376:. Indiana University Press. pp.
3220:, in addition to the above mentioned
3044:, but with wetter and drier seasons.
906:A map of caenagnathid discoveries in
116:A diagram of all known material from
7:
4094:Mallon, Jordan C.; S, Jason (2012).
2768:is substantially larger than either
1232:, and therefore probably belongs to
535:before being given its own genus in
3434:Timeline of oviraptorosaur research
2416:
2366:
2359:
2330:
2275:
2268:
2239:
2184:
2177:
2150:
2121:
2092:
2062:
2032:
2005:
1976:
1966:
1745:
1716:
1653:
1606:
1577:
1548:
1518:
1488:
1461:
1432:
1422:
1178:One of the metatarsals referred to
824:. Some specimens were referred to
796:. In his monograph, Sues referred
576:. This locality was a part of the
4511:Journal of Vertebrate Paleontology
4170:Canadian Journal of Earth Sciences
4131:Canadian Journal of Earth Sciences
3878:Canadian Journal of Earth Sciences
3652:Journal of Vertebrate Paleontology
3610:Canadian Journal of Earth Sciences
3566:Canadian Journal of Earth Sciences
3496:Funston, Gregory (July 27, 2020).
3010:, marks the top of the formation.
1947:. The controversial Asian genera
1932:caenagnathids. He also recovered
808:, and he also regarded the genera
672:Canadian Journal of Earth Sciences
14:
5441:Campanian genus first appearances
4168:(Dinosauria: Oviraptorosauria)".
2878:. This is unlikely to be due to
2681:One of the specimens referred to
580:, which is now recognized as the
5308:
5307:
5284:
5279:
3414:List of North American dinosaurs
3200:is also known to have preyed on
2914:could be considered prey items.
2806:remains preserved in its stomach
2428:
2342:
2304:
2251:
2196:
2133:
2104:
2072:
2042:
2015:
1988:
1728:
1635:
1589:
1560:
1528:
1498:
1471:
1444:
882:to the new genus as the species
828:and others to the newly erected
789:was a distinct and valid genus.
718:
707:
405:
129:
4271:Fowler, Denver Warwick (2017).
3006:interbedded with mudstones and
493:" is Latin for "elegant". The
3672:10.1080/02724634.1997.10011018
3646:Sues, Hans-Dieter (1997). "On
3392:2023 in archosaur paleontology
3387:2020 in archosaur paleontology
2824:individuals. The legs of the
596:and was given the designation
1:
5461:Fossil taxa described in 2020
4531:10.1080/02724634.2019.1649681
4020:10.1016/j.cretres.2020.104521
3120:in such well-known genera as
3048:were apparently the dominant
1187:New material assigned in 2020
1143:elements. They also erected
1097:Referred material (1989-2013)
939:Mongolian Academy of Sciences
4298:10.1371/journal.pone.0188426
4116:10.1016/j.palaeo.2012.06.024
3317:. There were also abundant
3029:conditions over time as the
485:for "fleet-footed", and the
5446:Campanian genus extinctions
3080:An artistic rendition of a
2722:Diet and niche partitioning
747:was distinguished from the
337:(Parks, 1933) Russell, 1972
301:(Parks, 1933) Funston, 2020
5482:
3942:Royal Society Open Science
3855:Doklady Akademii Nauk SSSR
3285:. The recently-described
2906:individuals of both major
1147:(the anterior half of two
390:
16:Genus of reptiles (fossil)
5466:Monotypic dinosaur genera
5297:
5277:
5079:
4912:
4802:
4798:
4783:
4764:
4714:
4687:
4660:
4647:
4586:
4366:Eberth, David A. (2005).
4166:Chirostenotes pergracilis
3874:Chirostenotes pergracilis
3822:Canadian Field-Naturalist
3785:American Museum Novitates
3311:and the widespread genus
3255:taxa, although only one,
3184:, which coexisted in the
3104:including the well-known
2443:
2421:
2414:
2388:
2371:
2364:
2357:
2335:
2328:
2297:
2280:
2273:
2266:
2244:
2237:
2211:
2189:
2182:
2175:
2155:
2148:
2126:
2119:
2097:
2090:
2067:
2060:
2037:
2030:
2010:
2003:
1981:
1974:
1767:
1750:
1743:
1721:
1714:
1697:
1680:
1658:
1651:
1628:
1611:
1604:
1582:
1575:
1553:
1546:
1523:
1516:
1493:
1486:
1466:
1459:
1437:
1430:
923:Canadian Museum of Nature
880:Chirostenotes pergracilis
621:because of the lack of a
315:
308:
289:
284:
265:
258:
126:Scientific classification
124:
115:
106:
23:
4408:Dinosaur Provincial Park
4403:"Campanian palynomorphs"
4373:Dinosaur Provincial Park
3816:Sternberg, C.M. (1932).
2935:Dinosaur Provincial Park
1128:to refer the species to
1038:, he identifies several
917:material known from the
908:Dinosaur Provincial Park
391:Not to be confused with
5456:Paleontology in Alberta
3186:Dinosaur Park Formation
3094:Dinosaur Park Formation
3031:Western Interior Seaway
2959:Western Interior Seaway
2954:Dinosaur Park Formation
2947:Dinosaur Park Formation
2728:Dinosaur Park Formation
2663:Dinosaur Park Formation
2600:Dinosaur Park Formation
1886:Dinosaur Park Formation
1360:and a clade containing
984:Dinosaur Park Formation
919:Dinosaur Park Formation
890:from the type species,
737:was originally referred
698:Dinosaur Park Formation
592:, was reposited at the
582:Dinosaur Park Formation
477:. The generic name of
457:Dinosaur Park Formation
378:(Parks, 1933) Longrich
4239:10.1126/sciadv.adi0505
4190:10.1139/cjes-2013-0186
3401:Two Medicine Formation
3397:Judith River Formation
3089:
2927:
2836:occasionally fed upon
2807:
2669:study of caenagnathid
2657:
2588:
1888:, Funston conducted a
1282:
1183:
1071:
910:
832:. Sues distinguished
759:fourth metatarsal bone
561:
5403:Paleobiology Database
5368:Leptorhynchos elegans
5338:Leptorhynchos elegans
4044:The Anatomical Record
3079:
2925:
2797:
2687:Leptorhynchos elegans
2652:
2586:
1754:Caenagnathus collinsi
1684:Leptorhynchos gaddisi
1662:Leptorhynchos elegans
1285:The first systematic
1280:
1241:tarsometatarsal bones
1177:
1137:Leptorhynchos elegans
1065:
1020:Histological analysis
941:. He also collected
935:University of Alberta
905:
898:Current understanding
884:Leptorhynchos elegans
623:fifth metatarsal bone
586:University of Toronto
570:University of Toronto
555:
4102:. 350–352: 124–138.
3762:10.3374/014.054.0204
3722:10.3374/014.054.0102
3345:mammal groups — the
2855:. Histology of the
2645:Growth and histology
1774:sp. (BMNH 2033 from
1124:, which is what led
1002:. In his review of
927:Royal Ontario Museum
878:, and so they moved
690:Royal Ontario Museum
611:Ornithomimus elegans
594:Royal Ontario Museum
543:Discovery and naming
271:Ornithomimus elegans
4844:Scansoriopterygidae
4523:2019JVPal..39E9681H
4455:10.7717/peerj.10720
4289:2017PLoSO..1288426F
4230:2023SciA....9I.505T
4182:2014CaJES..51..156F
4143:1993CaJES..30.2255C
4108:2012PPP...350..124M
4012:2020CrRes.11404521R
4000:Cretaceous Research
3962:10.1098/rsos.201184
3954:2020RSOS....701184F
3890:1988CaJES..25..972C
3664:1997JVPal..17..698S
3622:1989CaJES..26.1319C
3578:1972CaJES...9..375R
2579:Possible synonymity
1328:mandibular fenestra
1258:Royal Tyrell Museum
931:Royal Tyrell Museum
804:as the new species
663:a few years later.
34:~76.9 to 75.8 Ma -
3515:10.18435/vamp29362
3090:
3072:Contemporary fauna
3025:and influenced by
2928:
2808:
2658:
2591:In a study of the
2589:
1909:was not a natural
1283:
1184:
1072:
998:is a small-bodied
949:of the specimens.
911:
798:Elmisaurus elegans
745:Elmisaurus elegans
682:Elmisaurus elegans
562:
5418:
5417:
5390:Open Tree of Life
5330:Taxon identifiers
5321:
5320:
5293:
5292:
5275:
5274:
5271:
5270:
5267:
5266:
5066:
5065:
4888:
4887:
4872:Similicaudipteryx
4837:Protarchaeopteryx
4779:
4778:
4775:
4774:
4760:
4759:
4756:
4755:
4752:
4751:
4505:Cryodrakon boreas
4422:978-0-253-34595-0
4387:978-0-253-34595-0
4137:(10): 2255–2272.
3355:multituberculates
3245:Saurornitholestes
3021:that became more
2967:Bearpaw Formation
2575:
2574:
2566:
2565:
2557:
2556:
2548:
2547:
2539:
2538:
2530:
2529:
2521:
2520:
2512:
2511:
2503:
2502:
2494:
2493:
2485:
2484:
2476:
2475:
2467:
2466:
2458:
2457:
2403:
2402:
2317:
2316:
2226:
2225:
1880:In his review of
1877:
1876:
1868:
1867:
1859:
1858:
1850:
1849:
1841:
1840:
1832:
1831:
1823:
1822:
1814:
1813:
1805:
1804:
1796:
1795:
1787:
1786:
954:ontogenetic stage
645:Taxonomic history
578:Belly River Group
548:Initial discovery
501:has a convoluted
389:
388:
383:
368:
353:
338:
327:
302:
254:
5473:
5411:
5410:
5398:
5397:
5385:
5384:
5372:
5371:
5370:
5357:
5356:
5355:
5325:
5311:
5310:
5288:
5283:
5077:
5057:Epichirostenotes
4910:
4901:
4800:
4796:
4790:Oviraptorosauria
4785:
4768:Oviraptorosauria
4734:Oviraptorosauria
4712:
4685:
4658:
4649:
4642:
4641:
4588:
4580:Oviraptorosauria
4573:
4566:
4559:
4550:
4543:
4542:
4498:
4492:
4491:
4485:
4477:
4467:
4457:
4433:
4427:
4426:
4398:
4392:
4391:
4363:
4357:
4340:
4321:
4320:
4310:
4300:
4283:(11): e0188426.
4268:
4262:
4261:
4251:
4241:
4224:(49): eadi0505.
4218:Science Advances
4209:
4194:
4193:
4161:
4155:
4154:
4126:
4120:
4119:
4091:
4085:
4084:
4074:
4068:
4067:
4056:10.1002/ar.24205
4038:
4032:
4031:
3995:
3984:
3983:
3973:
3933:
3927:
3926:
3908:
3902:
3901:
3869:
3863:
3862:
3846:
3840:
3839:
3837:
3835:10.5962/p.339375
3813:
3807:
3806:
3796:
3780:
3774:
3773:
3745:
3734:
3733:
3701:
3676:
3675:
3643:
3634:
3633:
3616:(6): 1319–1324.
3601:
3590:
3589:
3561:
3552:
3551:
3539:
3528:
3527:
3517:
3493:
3304:nanhsiungchelyid
3210:ornithomimosaurs
3052:plants, with an
2918:Paleoenvironment
2872:caudal vertebrae
2432:
2417:
2392:Epichirostenotes
2367:
2360:
2346:
2331:
2308:
2276:
2269:
2255:
2240:
2200:
2185:
2178:
2151:
2137:
2122:
2108:
2093:
2076:
2063:
2046:
2033:
2019:
2006:
1992:
1977:
1971:Oviraptorosauria
1967:
1944:Epichirostenotes
1894:oviraptorosauria
1746:
1732:
1717:
1654:
1639:
1607:
1593:
1578:
1564:
1549:
1532:
1519:
1502:
1489:
1475:
1462:
1448:
1433:
1427:Oviraptorosauria
1423:
1330:, and a shallow
1117:Elmisaurus rarus
1114:similarities to
722:
711:
668:Philip J. Currie
627:Oviraptorosauria
437:
436:
433:
432:
429:
426:
423:
420:
417:
414:
411:
377:
362:
347:
336:
325:
300:
294:
268:
249:
242:
231:
226:
213:
134:
133:
111:
101:
38:
29:Temporal range:
21:
5481:
5480:
5476:
5475:
5474:
5472:
5471:
5470:
5421:
5420:
5419:
5414:
5406:
5401:
5393:
5388:
5380:
5375:
5366:
5365:
5360:
5351:
5350:
5345:
5332:
5322:
5317:
5289:
5263:
5170:
5119:
5062:
5014:
4987:
4982:Ojoraptorsaurus
4897:
4895:Caenagnathoidea
4884:
4879:Xingtianosaurus
4850:
4792:
4771:
4748:
4728:
4701:
4674:
4668:Avemetatarsalia
4653:Avemetatarsalia
4643:
4593:
4592:
4582:
4577:
4547:
4546:
4517:(3): e1649681.
4500:
4499:
4495:
4478:
4435:
4434:
4430:
4423:
4400:
4399:
4395:
4388:
4365:
4364:
4360:
4341:
4324:
4270:
4269:
4265:
4211:
4210:
4197:
4163:
4162:
4158:
4151:10.1139/e93-196
4128:
4127:
4123:
4093:
4092:
4088:
4076:
4075:
4071:
4040:
4039:
4035:
3997:
3996:
3987:
3935:
3934:
3930:
3923:
3910:
3909:
3905:
3898:10.1139/e88-097
3871:
3870:
3866:
3848:
3847:
3843:
3815:
3814:
3810:
3782:
3781:
3777:
3747:
3746:
3737:
3703:
3702:
3679:
3645:
3644:
3637:
3630:10.1139/e89-111
3603:
3602:
3593:
3586:10.1139/e72-031
3563:
3562:
3555:
3541:
3540:
3531:
3495:
3494:
3447:
3442:
3383:
3343:Late Cretaceous
3188:as well as the
3107:Parasaurolophus
3074:
2920:
2792:
2724:
2719:
2711:trabecular bone
2679:
2647:
2642:
2581:
2576:
2567:
2558:
2549:
2540:
2531:
2522:
2513:
2504:
2495:
2486:
2477:
2468:
2459:
2404:
2347:
2318:
2227:
2057:Caenagnathoidea
2047:
1878:
1869:
1860:
1851:
1842:
1833:
1824:
1815:
1806:
1797:
1788:
1649:Caenagnathinae
1513:Caenagnathoidea
1503:
1413:oviraptorosaurs
1332:surangular bone
1275:
1270:
1217:oviraptorosaurs
1205:sacral vertebra
1145:TMP 1992.36.290
1141:tarsometatarsal
1122:arctometatarsus
1110:. The overall
993:
947:ontogenetic age
900:
818:junior synonyms
742:
741:
740:
739:
738:
723:
714:
713:
712:
694:tarsometatarsus
647:
619:ornithomimosaur
550:
545:
454:Late Cretaceous
408:
404:
396:
324:
297:Citipes elegans
280:
274:
248:
240:
229:
224:
211:
128:
102:
100:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
36:
35:
33:
31:Late Cretaceous
27:
17:
12:
11:
5:
5479:
5477:
5469:
5468:
5463:
5458:
5453:
5451:Campanian life
5448:
5443:
5438:
5433:
5423:
5422:
5416:
5415:
5413:
5412:
5399:
5386:
5373:
5358:
5342:
5340:
5334:
5333:
5328:
5319:
5318:
5316:
5315:
5305:
5298:
5295:
5294:
5291:
5290:
5278:
5276:
5273:
5272:
5269:
5268:
5265:
5264:
5262:
5261:
5253:
5246:
5239:
5236:Machairasaurus
5232:
5225:
5218:
5211:
5204:
5196:
5189:
5180:
5178:
5172:
5171:
5169:
5168:
5160:
5153:
5145:
5137:
5129:
5127:
5121:
5120:
5118:
5117:
5110:
5103:
5095:
5088:
5080:
5074:
5068:
5067:
5064:
5063:
5061:
5060:
5053:
5046:
5043:Caenagnathasia
5039:
5032:
5024:
5022:
5020:Caenagnathinae
5016:
5015:
5013:
5012:
5005:
4997:
4995:
4989:
4988:
4986:
4985:
4978:
4970:
4963:
4956:
4949:
4942:
4935:
4928:
4921:
4913:
4907:
4905:Caenagnathidae
4898:
4893:
4890:
4889:
4886:
4885:
4883:
4882:
4875:
4868:
4860:
4858:
4852:
4851:
4849:
4848:
4840:
4833:
4830:Ningyuansaurus
4826:
4818:
4815:Incisivosaurus
4811:
4803:
4793:
4788:
4781:
4780:
4777:
4776:
4773:
4772:
4765:
4762:
4761:
4758:
4757:
4754:
4753:
4750:
4749:
4747:
4746:
4745:
4744:
4738:
4736:
4730:
4729:
4727:
4726:
4725:
4724:
4715:
4709:
4703:
4702:
4700:
4699:
4698:
4697:
4688:
4682:
4676:
4675:
4673:
4672:
4671:
4670:
4661:
4655:
4645:
4644:
4640:
4639:
4630:
4621:
4612:
4606:
4600:
4591:
4584:
4583:
4578:
4576:
4575:
4568:
4561:
4553:
4545:
4544:
4493:
4428:
4421:
4393:
4386:
4358:
4322:
4263:
4195:
4176:(2): 156–165.
4156:
4121:
4086:
4069:
4050:(4): 918–934.
4033:
3985:
3928:
3922:978-0691180311
3921:
3903:
3884:(7): 972–986.
3864:
3841:
3808:
3775:
3756:(2): 263–264.
3735:
3677:
3658:(4): 698–716.
3635:
3591:
3572:(4): 375–402.
3553:
3529:
3444:
3443:
3441:
3438:
3437:
3436:
3431:
3421:
3416:
3411:
3394:
3389:
3382:
3379:
3337:being known.
3323:helodermatidae
3276:Albertochampsa
3239:Latenivenatrix
3181:Daspletosaurus
3171:tyrannosaurids
3167:ornithischians
3159:Euoplocephalus
3154:ankylosauridae
3084:(right) and a
3082:Euoplocephalus
3073:
3070:
3036:westward. The
2919:
2916:
2791:
2788:
2723:
2720:
2718:
2715:
2685:(at the time,
2678:
2675:
2646:
2643:
2641:
2638:
2608:pubic peduncle
2580:
2577:
2573:
2572:
2569:
2568:
2564:
2563:
2560:
2559:
2555:
2554:
2551:
2550:
2546:
2545:
2542:
2541:
2537:
2536:
2533:
2532:
2528:
2527:
2524:
2523:
2519:
2518:
2515:
2514:
2510:
2509:
2506:
2505:
2501:
2500:
2497:
2496:
2492:
2491:
2488:
2487:
2483:
2482:
2479:
2478:
2474:
2473:
2470:
2469:
2465:
2464:
2461:
2460:
2456:
2455:
2452:
2451:
2442:
2439:
2438:
2435:
2434:
2420:
2415:
2413:
2410:
2409:
2406:
2405:
2401:
2400:
2397:
2396:
2387:
2384:
2383:
2380:
2379:
2375:Caenagnathasia
2370:
2365:
2363:
2358:
2356:
2353:
2352:
2349:
2348:
2341:
2334:
2329:
2327:
2324:
2323:
2320:
2319:
2315:
2314:
2311:
2310:
2296:
2293:
2292:
2289:
2288:
2279:
2274:
2272:
2267:
2265:
2262:
2261:
2258:
2257:
2243:
2238:
2236:
2233:
2232:
2229:
2228:
2224:
2223:
2220:
2219:
2210:
2207:
2206:
2203:
2202:
2188:
2183:
2181:
2176:
2174:
2172:Caenagnathinae
2168:
2167:
2164:
2163:
2154:
2149:
2147:
2144:
2143:
2140:
2139:
2125:
2120:
2118:
2115:
2114:
2111:
2110:
2096:
2091:
2089:
2087:Caenagnathidae
2083:
2082:
2079:
2078:
2066:
2061:
2059:
2053:
2052:
2049:
2048:
2041:
2036:
2031:
2029:
2026:
2025:
2022:
2021:
2009:
2004:
2002:
1999:
1998:
1995:
1994:
1985:Incisivosaurus
1980:
1975:
1973:
1965:
1935:Caenagnathasia
1875:
1874:
1871:
1870:
1866:
1865:
1862:
1861:
1857:
1856:
1853:
1852:
1848:
1847:
1844:
1843:
1839:
1838:
1835:
1834:
1830:
1829:
1826:
1825:
1821:
1820:
1817:
1816:
1812:
1811:
1808:
1807:
1803:
1802:
1799:
1798:
1794:
1793:
1790:
1789:
1785:
1784:
1781:
1780:
1766:
1763:
1762:
1759:
1758:
1749:
1744:
1742:
1739:
1738:
1735:
1734:
1720:
1715:
1713:
1710:
1709:
1706:
1705:
1696:
1693:
1692:
1689:
1688:
1679:
1676:
1675:
1672:
1671:
1657:
1652:
1650:
1646:
1645:
1642:
1641:
1627:
1624:
1623:
1620:
1619:
1615:Caenagnathasia
1610:
1605:
1603:
1600:
1599:
1596:
1595:
1581:
1576:
1574:
1571:
1570:
1567:
1566:
1552:
1547:
1545:
1543:Caenagnathidae
1539:
1538:
1535:
1534:
1522:
1517:
1515:
1509:
1508:
1505:
1504:
1497:
1492:
1487:
1485:
1482:
1481:
1478:
1477:
1465:
1460:
1458:
1455:
1454:
1451:
1450:
1441:Incisivosaurus
1436:
1431:
1429:
1421:
1274:
1271:
1269:
1268:Classification
1266:
1189:
1188:
1099:
1098:
1077:
1076:
1040:autapomorphies
992:
989:
899:
896:
838:C. pergracilis
826:C. pergracilis
816:as subjective
724:
717:
716:
715:
706:
705:
704:
703:
702:
655:junior synonym
651:Macrophalangia
646:
643:
574:Red Deer River
549:
546:
544:
541:
514:Macrophalangia
387:
386:
385:
384:
369:
363:(Parks, 1933)
354:
348:(Parks, 1933)
339:
332:Macrophalangia
328:
313:
312:
306:
305:
304:
303:
287:
286:
282:
281:
275:
263:
262:
256:
255:
238:
234:
233:
222:
218:
217:
215:Caenagnathidae
209:
205:
204:
199:
192:
191:
186:
179:
178:
173:
166:
165:
160:
156:
155:
150:
146:
145:
140:
136:
135:
122:
121:
113:
112:
104:
103:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
39:
28:
15:
13:
10:
9:
6:
4:
3:
2:
5478:
5467:
5464:
5462:
5459:
5457:
5454:
5452:
5449:
5447:
5444:
5442:
5439:
5437:
5434:
5432:
5431:Caenagnathids
5429:
5428:
5426:
5409:
5404:
5400:
5396:
5391:
5387:
5383:
5378:
5374:
5369:
5363:
5359:
5354:
5348:
5344:
5343:
5341:
5339:
5335:
5331:
5326:
5314:
5306:
5304:
5300:
5299:
5296:
5287:
5282:
5259:
5258:
5254:
5252:
5251:
5247:
5245:
5244:
5240:
5238:
5237:
5233:
5231:
5230:
5226:
5224:
5223:
5222:Jiangxisaurus
5219:
5217:
5216:
5212:
5210:
5209:
5205:
5202:
5201:
5200:Ganzhousaurus
5197:
5195:
5194:
5190:
5187:
5186:
5182:
5181:
5179:
5177:
5173:
5166:
5165:
5161:
5159:
5158:
5154:
5151:
5150:
5146:
5143:
5142:
5141:Corythoraptor
5138:
5136:
5135:
5131:
5130:
5128:
5126:
5125:Oviraptorinae
5122:
5116:
5115:
5111:
5109:
5108:
5104:
5101:
5100:
5096:
5094:
5093:
5089:
5087:
5086:
5082:
5081:
5078:
5075:
5073:
5072:Oviraptoridae
5069:
5059:
5058:
5054:
5052:
5051:
5047:
5045:
5044:
5040:
5038:
5037:
5033:
5031:
5030:
5026:
5025:
5023:
5021:
5017:
5011:
5010:
5006:
5004:
5003:
4999:
4998:
4996:
4994:
4990:
4984:
4983:
4979:
4976:
4975:
4971:
4969:
4968:
4964:
4962:
4961:
4957:
4955:
4954:
4953:Leptorhynchos
4950:
4948:
4947:
4946:Gigantoraptor
4943:
4941:
4940:
4936:
4934:
4933:
4932:Chirostenotes
4929:
4927:
4926:
4922:
4920:
4919:
4915:
4914:
4911:
4908:
4906:
4902:
4899:
4896:
4891:
4881:
4880:
4876:
4874:
4873:
4869:
4867:
4866:
4862:
4861:
4859:
4857:
4856:Caudipteridae
4853:
4846:
4845:
4841:
4839:
4838:
4834:
4832:
4831:
4827:
4824:
4823:
4819:
4817:
4816:
4812:
4810:
4809:
4805:
4804:
4801:
4797:
4794:
4791:
4786:
4782:
4770:
4769:
4763:
4742:
4741:
4740:
4739:
4737:
4735:
4731:
4723:
4719:
4718:
4717:
4716:
4713:
4710:
4708:
4704:
4696:
4692:
4691:
4690:
4689:
4686:
4683:
4681:
4677:
4669:
4665:
4664:
4663:
4662:
4659:
4656:
4654:
4650:
4646:
4638:
4634:
4631:
4629:
4625:
4622:
4620:
4616:
4613:
4611:
4607:
4605:
4601:
4599:
4595:
4594:
4589:
4585:
4581:
4574:
4569:
4567:
4562:
4560:
4555:
4554:
4551:
4540:
4536:
4532:
4528:
4524:
4520:
4516:
4512:
4508:
4506:
4497:
4494:
4489:
4483:
4475:
4471:
4466:
4461:
4456:
4451:
4447:
4443:
4439:
4432:
4429:
4424:
4418:
4414:
4410:
4409:
4404:
4397:
4394:
4389:
4383:
4379:
4375:
4374:
4369:
4368:"The Geology"
4362:
4359:
4355:
4354:0-253-34595-2
4351:
4347:
4346:
4339:
4337:
4335:
4333:
4331:
4329:
4327:
4323:
4318:
4314:
4309:
4304:
4299:
4294:
4290:
4286:
4282:
4278:
4274:
4267:
4264:
4259:
4255:
4250:
4245:
4240:
4235:
4231:
4227:
4223:
4219:
4215:
4208:
4206:
4204:
4202:
4200:
4196:
4191:
4187:
4183:
4179:
4175:
4171:
4167:
4160:
4157:
4152:
4148:
4144:
4140:
4136:
4132:
4125:
4122:
4117:
4113:
4109:
4105:
4101:
4097:
4090:
4087:
4082:
4081:
4073:
4070:
4065:
4061:
4057:
4053:
4049:
4045:
4037:
4034:
4029:
4025:
4021:
4017:
4013:
4009:
4005:
4001:
3994:
3992:
3990:
3986:
3981:
3977:
3972:
3967:
3963:
3959:
3955:
3951:
3947:
3943:
3939:
3932:
3929:
3924:
3918:
3914:
3907:
3904:
3899:
3895:
3891:
3887:
3883:
3879:
3875:
3868:
3865:
3861:(3): 685−688.
3860:
3856:
3852:
3845:
3842:
3836:
3831:
3828:(5): 99–105.
3827:
3823:
3819:
3812:
3809:
3804:
3800:
3795:
3790:
3787:(144): 1−12.
3786:
3779:
3776:
3771:
3767:
3763:
3759:
3755:
3751:
3744:
3742:
3740:
3736:
3731:
3727:
3723:
3719:
3715:
3711:
3707:
3700:
3698:
3696:
3694:
3692:
3690:
3688:
3686:
3684:
3682:
3678:
3673:
3669:
3665:
3661:
3657:
3653:
3649:
3648:Chirostenotes
3642:
3640:
3636:
3631:
3627:
3623:
3619:
3615:
3611:
3607:
3600:
3598:
3596:
3592:
3587:
3583:
3579:
3575:
3571:
3567:
3560:
3558:
3554:
3549:
3545:
3538:
3536:
3534:
3530:
3525:
3521:
3516:
3511:
3507:
3503:
3499:
3492:
3490:
3488:
3486:
3484:
3482:
3480:
3478:
3476:
3474:
3472:
3470:
3468:
3466:
3464:
3462:
3460:
3458:
3456:
3454:
3452:
3450:
3446:
3439:
3435:
3432:
3429:
3425:
3422:
3420:
3417:
3415:
3412:
3410:
3406:
3402:
3398:
3395:
3393:
3390:
3388:
3385:
3384:
3380:
3378:
3376:
3372:
3368:
3364:
3360:
3359:carpet sharks
3356:
3352:
3348:
3344:
3340:
3336:
3332:
3328:
3324:
3320:
3316:
3315:
3310:
3309:
3305:
3299:
3297:
3293:
3292:
3288:
3284:
3283:
3278:
3277:
3273:
3269:
3268:
3267:Champsosaurus
3264:
3260:
3259:
3254:
3249:
3247:
3246:
3241:
3240:
3235:
3234:
3233:Dromaeosaurus
3229:
3228:
3223:
3222:caenagnathids
3219:
3215:
3211:
3207:
3203:
3199:
3196:formations.
3195:
3191:
3187:
3183:
3182:
3177:
3176:
3172:
3168:
3163:
3161:
3160:
3155:
3152:, as well as
3151:
3150:
3145:
3141:
3137:
3136:
3131:
3130:
3125:
3124:
3123:Styracosaurus
3119:
3115:
3114:
3113:Corythosaurus
3109:
3108:
3103:
3099:
3095:
3087:
3086:Styracosaurus
3083:
3078:
3071:
3069:
3067:
3063:
3059:
3055:
3051:
3047:
3043:
3039:
3035:
3032:
3028:
3024:
3020:
3016:
3011:
3009:
3005:
3002:
2998:
2994:
2990:
2986:
2982:
2979:
2975:
2970:
2968:
2964:
2960:
2955:
2950:
2948:
2944:
2940:
2936:
2932:
2924:
2917:
2915:
2913:
2909:
2905:
2901:
2895:
2893:
2889:
2885:
2881:
2877:
2873:
2869:
2866:Notably, the
2864:
2862:
2858:
2854:
2850:
2846:
2841:
2839:
2835:
2831:
2827:
2823:
2819:
2818:
2813:
2805:
2801:
2796:
2789:
2787:
2785:
2784:Chirostenotes
2781:
2780:
2775:
2774:Chirostenotes
2771:
2767:
2766:
2761:
2758:
2754:
2750:
2746:
2745:
2744:Chirostenotes
2740:
2739:
2732:
2729:
2721:
2716:
2714:
2712:
2708:
2704:
2700:
2696:
2692:
2688:
2684:
2676:
2674:
2672:
2668:
2664:
2656:
2651:
2644:
2639:
2637:
2635:
2634:Leptorhynchos
2631:
2630:
2629:Leptorhynchos
2625:
2621:
2615:
2613:
2609:
2605:
2601:
2597:
2594:
2585:
2578:
2571:
2570:
2562:
2561:
2553:
2552:
2544:
2543:
2535:
2534:
2526:
2525:
2517:
2516:
2508:
2507:
2499:
2498:
2490:
2489:
2481:
2480:
2472:
2471:
2463:
2462:
2454:
2453:
2450:
2449:
2448:
2441:
2440:
2437:
2436:
2433:
2431:
2427:
2426:
2419:
2418:
2412:
2411:
2408:
2407:
2399:
2398:
2395:
2394:
2393:
2386:
2385:
2382:
2381:
2378:
2377:
2376:
2369:
2368:
2362:
2361:
2355:
2354:
2351:
2350:
2345:
2340:
2339:
2333:
2332:
2326:
2325:
2322:
2321:
2313:
2312:
2309:
2307:
2303:
2302:
2295:
2294:
2291:
2290:
2287:
2286:
2285:
2278:
2277:
2271:
2270:
2264:
2263:
2260:
2259:
2256:
2254:
2250:
2249:
2242:
2241:
2235:
2234:
2231:
2230:
2222:
2221:
2218:
2217:
2216:
2209:
2208:
2205:
2204:
2201:
2199:
2195:
2194:
2193:Chirostenotes
2187:
2186:
2180:
2179:
2173:
2170:
2169:
2166:
2165:
2162:
2161:
2160:
2153:
2152:
2146:
2145:
2142:
2141:
2138:
2136:
2132:
2131:
2130:Gigantoraptor
2124:
2123:
2117:
2116:
2113:
2112:
2109:
2107:
2103:
2102:
2095:
2094:
2088:
2085:
2084:
2081:
2080:
2077:
2075:
2071:
2070:Oviraptoridae
2065:
2064:
2058:
2055:
2054:
2051:
2050:
2045:
2040:
2035:
2034:
2028:
2027:
2024:
2023:
2020:
2018:
2014:
2013:Caudipteridae
2008:
2007:
2001:
2000:
1997:
1996:
1993:
1991:
1987:
1986:
1979:
1978:
1972:
1969:
1968:
1964:
1962:
1961:oviraptoridae
1958:
1957:
1952:
1951:
1946:
1945:
1941:
1937:
1936:
1931:
1927:
1926:
1921:
1920:
1919:Chirostenotes
1915:
1912:
1908:
1903:
1901:
1900:
1895:
1891:
1887:
1883:
1882:caenagnathids
1873:
1872:
1864:
1863:
1855:
1854:
1846:
1845:
1837:
1836:
1828:
1827:
1819:
1818:
1810:
1809:
1801:
1800:
1792:
1791:
1783:
1782:
1779:
1777:
1773:
1772:
1765:
1764:
1761:
1760:
1757:
1756:
1755:
1748:
1747:
1741:
1740:
1737:
1736:
1733:
1731:
1727:
1726:
1725:Chirostenotes
1719:
1718:
1712:
1711:
1708:
1707:
1704:
1703:
1702:
1695:
1694:
1691:
1690:
1687:
1686:
1685:
1678:
1677:
1674:
1673:
1670:
1668:
1664:
1663:
1656:
1655:
1648:
1647:
1644:
1643:
1640:
1638:
1634:
1633:
1626:
1625:
1622:
1621:
1618:
1617:
1616:
1609:
1608:
1602:
1601:
1598:
1597:
1594:
1592:
1588:
1587:
1586:Gigantoraptor
1580:
1579:
1573:
1572:
1569:
1568:
1565:
1563:
1559:
1558:
1551:
1550:
1544:
1541:
1540:
1537:
1536:
1533:
1531:
1527:
1526:Oviraptoridae
1521:
1520:
1514:
1511:
1510:
1507:
1506:
1501:
1496:
1491:
1490:
1484:
1483:
1480:
1479:
1476:
1474:
1470:
1469:Caudipteridae
1464:
1463:
1457:
1456:
1453:
1452:
1449:
1447:
1443:
1442:
1435:
1434:
1428:
1425:
1424:
1420:
1416:
1414:
1410:
1409:
1408:Edmontosaurus
1404:
1403:
1398:
1394:
1390:
1386:
1381:
1379:
1378:Leptorhynchos
1375:
1371:
1370:
1369:Chirostenotes
1365:
1364:
1359:
1358:
1353:
1349:
1348:Leptorhynchos
1345:
1344:
1339:
1338:
1337:Gigantoraptor
1333:
1329:
1325:
1321:
1316:
1314:
1313:
1308:
1307:
1302:
1298:
1297:Leptorhynchos
1294:
1293:
1292:Leptorhynchos
1288:
1279:
1272:
1267:
1265:
1263:
1262:Chirostenotes
1259:
1254:
1252:
1251:
1246:
1242:
1237:
1235:
1231:
1230:Chirostenotes
1227:
1222:
1218:
1214:
1210:
1206:
1201:
1198:
1194:
1186:
1185:
1181:
1176:
1172:
1170:
1166:
1165:Chirostenotes
1162:
1158:
1154:
1150:
1146:
1142:
1138:
1133:
1131:
1127:
1126:Philip Currie
1123:
1119:
1118:
1113:
1112:morphological
1109:
1104:
1096:
1095:
1094:
1092:
1091:
1086:
1082:
1074:
1073:
1070:— ROM 781 (C)
1069:
1064:
1060:
1058:
1057:
1056:Leptorhynchos
1052:
1051:
1046:
1041:
1037:
1032:
1029:
1025:
1021:
1017:
1016:
1015:Chirostenotes
1011:
1010:
1005:
1004:Dinosaur Park
1001:
997:
990:
988:
985:
980:
978:
974:
972:
971:Leptorhynchos
967:
966:
965:Chirostenotes
961:
960:
955:
950:
948:
944:
940:
936:
932:
928:
924:
920:
916:
909:
904:
897:
895:
893:
889:
885:
881:
877:
876:Leptorhynchos
873:
872:Chirostenotes
869:
865:
864:
859:
858:
857:Leptorhynchos
852:
850:
847:
843:
842:Chirostenotes
839:
835:
831:
827:
823:
822:Chirostenotes
819:
815:
811:
807:
803:
802:Chirostenotes
800:to the genus
799:
795:
794:Chirostenotes
790:
788:
787:
782:
781:
776:
775:
770:
769:
768:Chirostenotes
764:
760:
756:
755:
750:
746:
736:
732:
728:
727:Chirostenotes
721:
710:
701:
699:
695:
691:
687:
683:
679:
678:
673:
669:
664:
662:
661:
660:Chirostenotes
656:
652:
644:
642:
640:
639:
638:Chirostenotes
634:
633:
628:
624:
620:
616:
612:
608:
607:
601:
599:
595:
591:
587:
583:
579:
575:
571:
567:
559:
554:
547:
542:
540:
538:
534:
533:
532:Leptorhynchos
528:
527:
526:Chirostenotes
522:
521:
516:
515:
510:
509:
504:
500:
496:
495:type specimen
492:
488:
484:
480:
476:
475:
470:
466:
462:
458:
455:
451:
448:
444:
441:
435:
402:
401:
394:
381:
376:
374:
373:Leptorhynchos
370:
366:
361:
359:
358:Chirostenotes
355:
351:
346:
344:
340:
335:
333:
329:
323:
321:
317:
316:
314:
311:
307:
299:
298:
291:
290:
288:
283:
278:
273:
272:
264:
261:
257:
253:
247:
246:
239:
236:
235:
232:
223:
220:
219:
216:
210:
207:
206:
203:
200:
197:
194:
193:
190:
187:
184:
181:
180:
177:
174:
171:
168:
167:
164:
161:
158:
157:
154:
151:
148:
147:
144:
141:
138:
137:
132:
127:
123:
119:
114:
110:
105:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
32:
26:
22:
19:
5337:
5255:
5248:
5241:
5234:
5227:
5220:
5213:
5206:
5198:
5193:Conchoraptor
5191:
5183:
5162:
5155:
5149:Huanansaurus
5147:
5139:
5132:
5112:
5107:Tongtianlong
5105:
5097:
5090:
5083:
5055:
5050:Caenagnathus
5048:
5041:
5034:
5027:
5007:
5001:
5000:
4993:Elmisaurinae
4980:
4972:
4967:Microvenator
4965:
4958:
4951:
4944:
4937:
4930:
4923:
4916:
4877:
4870:
4863:
4842:
4835:
4828:
4820:
4813:
4806:
4766:
4637:Pennaraptora
4632:
4623:
4614:
4514:
4510:
4504:
4496:
4482:cite journal
4445:
4441:
4431:
4407:
4396:
4372:
4361:
4343:
4280:
4276:
4266:
4221:
4217:
4173:
4169:
4165:
4159:
4134:
4130:
4124:
4099:
4089:
4079:
4072:
4047:
4043:
4036:
4003:
3999:
3945:
3941:
3931:
3912:
3906:
3881:
3877:
3873:
3867:
3858:
3854:
3844:
3825:
3821:
3811:
3784:
3778:
3753:
3749:
3713:
3709:
3655:
3651:
3647:
3613:
3609:
3605:
3569:
3565:
3547:
3543:
3505:
3501:
3351:metatherians
3335:xenosauridae
3312:
3306:
3300:
3289:
3282:Leidyosuchus
3280:
3274:
3272:crocodilians
3265:
3263:choristodere
3258:Fluvionectes
3256:
3250:
3243:
3237:
3231:
3227:Paronychodon
3225:
3218:dromaeosaurs
3201:
3197:
3194:Two Medicine
3190:Judith River
3179:
3173:
3164:
3157:
3147:
3144:nodosauridae
3135:Centrosaurus
3133:
3129:Chasmosaurus
3127:
3121:
3111:
3105:
3102:hadrosaurids
3091:
3085:
3081:
3034:transgressed
3012:
2971:
2951:
2930:
2929:
2911:
2903:
2899:
2896:
2891:
2887:
2883:
2875:
2867:
2865:
2860:
2856:
2852:
2848:
2842:
2837:
2833:
2829:
2825:
2821:
2815:
2809:
2803:
2799:
2783:
2777:
2773:
2769:
2765:Caenagnathus
2763:
2762:
2757:cursoriality
2752:
2748:
2742:
2738:Caenagnathus
2736:
2733:
2725:
2717:Paleoecology
2686:
2682:
2680:
2667:histological
2659:
2654:
2640:Paleobiology
2633:
2627:
2623:
2616:
2603:
2590:
2447:Caenagnathus
2445:
2444:
2423:
2422:
2390:
2389:
2373:
2372:
2336:
2299:
2298:
2283:
2282:
2281:
2246:
2245:
2213:
2212:
2191:
2190:
2157:
2156:
2128:
2127:
2101:Microvenator
2099:
2098:
2068:
2011:
1983:
1982:
1954:
1948:
1942:
1933:
1923:
1917:
1911:monophyletic
1907:elmisaurinae
1904:
1897:
1892:analysis of
1890:phylogenetic
1879:
1771:Caenagnathus
1769:
1768:
1752:
1751:
1723:
1722:
1699:
1698:
1682:
1681:
1666:
1661:
1660:
1659:
1630:
1629:
1613:
1612:
1584:
1583:
1557:Microvenator
1555:
1554:
1524:
1467:
1439:
1438:
1417:
1406:
1400:
1385:caenagnathid
1382:
1377:
1374:Caenagnathus
1373:
1367:
1363:Caenagnathus
1361:
1355:
1347:
1343:Microvenator
1341:
1335:
1317:
1310:
1304:
1300:
1296:
1290:
1287:phylogenetic
1284:
1261:
1255:
1248:
1245:histological
1238:
1233:
1229:
1226:Caenagnathus
1225:
1202:
1192:
1190:
1179:
1168:
1164:
1160:
1156:
1144:
1136:
1134:
1129:
1115:
1100:
1090:Ornithomimus
1088:
1084:
1078:
1067:
1054:
1048:
1044:
1035:
1033:
1027:
1013:
1009:Caenagnathus
1007:
1000:caenagnathid
995:
994:
981:
976:
969:
963:
959:Caenagnathus
957:
951:
943:histological
915:caenagnathid
912:
891:
887:
883:
879:
875:
871:
868:Caenagnathus
867:
861:
855:
853:
841:
837:
833:
829:
825:
821:
813:
810:Caenagnathus
809:
805:
801:
797:
793:
791:
784:
778:
774:Caenagnathus
772:
766:
763:caenagnathid
752:
749:type species
744:
743:
734:
730:
726:
681:
675:
665:
658:
650:
648:
636:
630:
610:
606:Ornithomimus
604:
602:
597:
565:
563:
557:
530:
524:
518:
512:
508:Ornithomimus
506:
498:
490:
478:
473:
472:
469:type species
447:caenagnathid
399:
398:
397:
379:
371:
356:
341:
330:
320:Ornithomimus
318:
296:
295:
270:
269:
260:Type species
244:
243:
228:Elmisaurinae
195:
182:
169:
117:
24:
18:
5362:Wikispecies
5243:Nemegtomaia
5176:Heyuanninae
5036:Apatoraptor
4865:Caudipteryx
4722:Maniraptora
4707:Maniraptora
3508:: 105–153.
3214:troodontids
3198:Gorgosaurus
3175:Gorgosaurus
3140:Ankylosaurs
3118:ceratopsids
3066:angiosperms
3019:floodplains
2978:crossbedded
2908:ontogenetic
2904:Gorgosaurus
2900:Gorgosaurus
2884:Gorgosaurus
2876:Gorgosaurus
2861:Gorgosaurus
2853:Gorgosaurus
2849:Gorgosaurus
2834:Gorgosaurus
2830:Gorgosaurus
2817:Gorgosaurus
2800:Gorgosaurus
2798:A juvenile
2338:Apatoraptor
1925:Apatoraptor
1402:Triceratops
1306:Nemegtomaia
1221:pleurocoels
1024:metatarsals
991:Description
733:, to which
615:metatarsals
590:metatarsals
326:Parks, 1933
221:Subfamily:
5425:Categories
5301:See also:
5257:Shixinggia
5208:Gobiraptor
5085:Luoyanggia
5009:Elmisaurus
4939:Eoneophron
4925:Beibeilong
4918:Anomalipes
4743:see below↓
4619:Dinosauria
4610:Sauropsida
4448:: e10720.
3606:Elmisaurus
3440:References
3375:paddlefish
3347:eutherians
3291:Cryodrakon
3287:azhdarchid
3253:plesiosaur
3208:including
3156:, such as
3149:Edmontonia
3146:, such as
3062:tree ferns
3054:understory
3008:siltstones
2989:floodplain
2981:sandstones
2707:Taphonomic
2596:morphology
2301:Elmisaurus
2159:Anomalipes
2039:Avimimidae
1956:Anomalipes
1940:Laramidian
1930:Laramidian
1776:Hell Creek
1632:Elmisaurus
1495:Avimimidae
1301:L. elegans
1250:Elmisaurus
1169:L. elegans
1161:Elmisaurus
1157:L. elegans
1130:Elmisaurus
1108:taphonomic
1050:Elmisaurus
937:, and the
892:L. gaddisi
888:L. elegans
834:C. elegans
830:C. elegans
814:Elmisaurus
806:C. elegans
786:Elmisaurus
780:Elmisaurus
731:Elmisaurus
686:metatarsus
677:Elmisaurus
520:Elmisaurus
474:C. elegans
343:Elmisaurus
189:Saurischia
176:Dinosauria
5353:Q76824533
5215:Heyuannia
5164:Rinchenia
5157:Oviraptor
5092:Nankangia
4960:Hagryphus
4695:Theropoda
4680:Theropoda
4628:Theropoda
4596:Kingdom:
4539:203406859
4028:219745025
3851:theropods
3794:2246/3223
3770:128898931
3730:128444961
3716:: 23–49.
3524:2292-1389
3428:Judithian
3371:sturgeons
3363:stingrays
3327:varanidae
3308:Basilemys
3298:remains.
3296:pterosaur
3206:theropods
3169:were the
2997:bentonite
2993:mudstones
2880:digestion
2790:Predation
2699:ischiatic
2677:Pathology
2671:dentaries
2215:Hagryphus
1884:from the
1701:Hagryphus
1357:Hagryphus
1320:monophyly
1273:Phylogeny
1151:) as the
1149:dentaries
863:Hagryphus
849:Laramidia
846:Campanian
632:Oviraptor
560:was found
503:taxonomic
489:epithet "
452:from the
250:Funston,
230: (?)
202:Theropoda
149:Kingdom:
143:Eukaryota
37:Campanian
5347:Wikidata
5313:Category
5303:Timeline
5134:Citipati
5099:Nomingia
4974:Nomingia
4808:Avimimus
4604:Chordata
4602:Phylum:
4598:Animalia
4474:33614274
4317:29166406
4277:PLOS ONE
4258:38064561
4249:10846869
4064:31270950
3980:33204472
3803:40272928
3381:See also
3270:and the
3098:dinosaur
3046:Conifers
3001:low-rank
2985:overbank
2248:Nomingia
1950:Nomingia
1389:mandible
1352:polytomy
1312:Nomingia
1197:paratype
1153:paratype
1081:holotype
1075:Holotype
754:E. rarus
487:specific
450:theropod
438:) is an
393:Citipati
310:Synonyms
285:Species
208:Family:
163:Chordata
159:Phylum:
153:Animalia
139:Domain:
5395:4126519
5382:8502277
5002:Citipes
4608:Class:
4519:Bibcode
4465:7882142
4308:5699823
4285:Bibcode
4226:Bibcode
4178:Bibcode
4139:Bibcode
4104:Bibcode
4008:Bibcode
3971:7657903
3950:Bibcode
3886:Bibcode
3660:Bibcode
3618:Bibcode
3574:Bibcode
3550:: 4–44.
3409:Montana
3405:Alberta
3339:Mammals
3331:teiidae
3319:lizards
3202:Citipes
3038:climate
2974:channel
2965:of the
2939:Alberta
2931:Citipes
2912:Citipes
2892:Citipes
2888:Citipes
2868:Citipes
2857:Citipes
2847:of the
2838:Citipes
2826:Citipes
2822:Citipes
2804:Citipes
2770:Citipes
2753:Citipes
2749:Citipes
2683:Citipes
2655:Citipes
2624:Citipes
2604:Citipes
2284:Citipes
1667:Citipes
1665:(later
1324:dentary
1234:Citipes
1213:centrum
1193:Citipes
1180:Citipes
1085:Citipes
1068:Citipes
1045:Citipes
1036:Citipes
1028:Citipes
1022:of two
996:Citipes
977:Citipes
973:elegans
765:genera
735:Citipes
598:ROM 781
566:Citipes
558:Citipes
499:Citipes
491:elegans
479:Citipes
461:Alberta
440:extinct
400:Citipes
375:elegans
360:elegans
345:elegans
334:elegans
322:elegans
245:Citipes
237:Genus:
118:Citipes
25:Citipes
5408:266223
5250:Oksoko
5114:Yulong
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4413:101–30
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3353:, and
3333:, and
3314:Adocus
3242:, and
3216:, and
3132:, and
3064:, and
3050:canopy
3027:marine
3023:swampy
3015:rivers
2963:shales
2943:Canada
2747:, and
2593:pelvic
1899:Oksoko
933:, the
929:, the
925:, the
870:, and
777:, and
529:, and
465:Canada
382:, 2013
380:et al.
367:, 1997
352:, 1989
350:Currie
279:, 1933
5229:Khaan
5185:Banji
4633:Clade
4624:Clade
4615:Clade
4535:S2CID
4442:PeerJ
4378:54–82
4024:S2CID
3766:S2CID
3726:S2CID
3058:ferns
3042:frost
2845:femur
2802:with
2703:fossa
2695:pubic
2691:ilium
2620:sacra
2612:ilium
1914:clade
1397:birds
1393:manus
1354:with
1103:taxon
836:from
483:Latin
443:genus
277:Parks
196:Clade
183:Clade
170:Clade
5377:GBIF
5029:Anzu
4720:see
4693:see
4666:see
4488:link
4470:PMID
4417:ISBN
4382:ISBN
4350:ISBN
4313:PMID
4254:PMID
4060:PMID
3976:PMID
3917:ISBN
3799:OCLC
3520:ISSN
3407:and
3399:and
3279:and
3192:and
3178:and
3110:and
3092:The
3017:and
3004:coal
2987:and
2952:The
2843:The
2812:2023
2779:Anzu
2726:The
2697:and
2425:Anzu
1953:and
1922:and
1405:and
1391:and
1340:and
1318:The
1309:and
1209:ilia
1163:and
1079:The
1053:and
1012:and
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812:and
729:and
635:and
537:2020
365:Sues
252:2020
42:PreꞒ
4822:Kol
4527:doi
4460:PMC
4450:doi
4303:PMC
4293:doi
4244:PMC
4234:doi
4186:doi
4147:doi
4112:doi
4052:doi
4048:303
4016:doi
4004:114
3966:PMC
3958:doi
3894:doi
3859:226
3830:doi
3789:hdl
3758:doi
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