727:
nasal region are eight caputegulae that are ordered in a symmetrical pattern, but are separated by a row of pyramid-like caputegulae. Large, hexagonal caputegulae form a sagittal midline row which terminate in an apex towards the rear and towards the sides of sagittal midline row is a row of both polygonal and pyramid-shaped caputegulae. The side projecting nasal caputegulum forms a flaring nostril in side view. Positioned on the sides of each prefrontal is a subtriangular caputegulum with a keeled apex, which is anterior to the anterior-most supraorbital boss. A caputegulum that has a hexagonal base and a bulbous shape is surrounded by six smaller caputegulae towards the sides and front, which are similar to the nasal caputegulae but are different to the large, posterior caputegulum. The nuchal shelf has little rugosity and three, poorly preserved nuchal caputegulae, which vary between subrounded to elongate polygonal, are present on the posterior-most portion. The circumorbital complex consists of a supraorbital horn, a lacrimal caputegulum, a jugal osteoderm, and the thickened rim along the posterior margin of the orbital. The postorbital horn is formed by the fused anterior and posterior supraorbital caputegulae. Due to the damaged squamosal horns and more vertically oriented quadratojugal horns, it is unknown if
978:
distinct morphologies are known which include a “pup-tent” shaped osteoderm with keels that are S-twisted and have a backswept apex (type 1), triangular osteoderms that are compressed dorsoventrally (type 2), and flat-based osteoderms that have a keel that is off-centred and longitudinal (type 5). Type 1 osteoderms are represent by 5 mostly incomplete osteoderms which are asymmetrical and have rounded apexes. These osteoderms would have been positioned on the thoracic or pectoral region of the body. The surface texture of these type 1 osteoderms vary as some are minimally rugose while others are rugose and with densely distributed, shallow pitting. Type 2 osteoderms are represented by 8 partial osteoderms which all vary in size and have a surface texture which is slightly rugose with minimal pitting. The type 2 osteoderms have a morphology which is typically seen in osteoderms that are positioned along the sides of the thorax and tail, as seen in specimens of
4159:
604:
736:
719:
841:
distal caudal vertebrae in the proximal series as they are more round. The proximal ribs have dorsal surfaces that expand horizontally to form a T- or L-shaped cross-section, with three proximal ribs having an L-shaped cross-section and arch towards the underside. The proximal ribs with a T-shaped cross section have a wide horizontal expansion on the upper side of both sides of the ribs and arch less sharply, unlike the L-shaped ribs, which suggests they were positioned more towards the front of the vertebral column.
1792:
612:
4154:
136:
644:
401:
118:
796:
4182:
962:
845:
1009:
908:
627:, a central large flat hexagonal osteoderm is present. The zone spanning the frontal bones and nasal bones is covered by symmetrically positioned, closely packed, pyramid-shaped and conical caputegulae. The nasal bones exhibit a distinct central row of conical caputegulae, symmetrically separated from the osteoderms above and to the sides of them. At the rear of the skull, the part of the
3081:
3069:
732:
underside of the supraorbital boss, while the jugal caputegulum is concave towards the underside and extends towards the sides which creates a small shelf. The quadrates are visible towards the sides. A subtriangular caputegulum with a blunt keel is present on the mandible and is shorter than that of other ankylosaurids.
623:, unique derived characters. The supraorbital bosses are massive in side view, forming a high back-swept ridge, also extending sideways over the eye socket, while encompassing the front top corner and the rear edge of the eye socket. The cheek horns are triangular, pointing almost vertically to below. On the
832:. The sutural contacts are nearly concealed between individual vertebrae, which may be an ontogenetic feature. The dorsosacral vertebrae have short ribs that are fused to the centrum and transverse processes. These ribs, along with the sacral ribs, form broad contact surfaces that articulate with the
973:
and posterior cervical half ring are preserved, but the anterior and posterior margins of both half rings are broken. Each cervical half ring is composed of six individual osteoderms, which are fused to the upper surfaces of the half rings and lack saw-tooth sutures. The loci present on the cervical
876:
has an enlarged deltopectoral crest which is soon followed by a thick humeral shaft. A large projecting process is present towards the back and underside of the humeral head, which is present in some other ankylosaurids but are expressed to a lesser extent. On the side of the deltopectoral crest is
726:
The external nares are ornamented with a supranasal caputegulum that is arranged along the premaxillary beak at the back and the sides. The caputegulum has a smooth surface texture, with some rugosity, and a tetrahedral-shaped lacrimal caputegulum is present near it. Along the dorsal surface of the
934:
of the handle interlocks with the adjacent chevron and are compressed in a way to which they look like elongated rods in lateral view. In addition, the chevrons are v-shaped structures that are fused onto the underside of the centra of the caudal vertebrae. Fused onto the two posterior-most caudal
731:
had squamosal horns and quadratojugal horns that exceeded the width of the postorbital horns. The postorbital horns are triangular towards the back, with a rugose and bulbous surface texture. The lacrimal caputegulum is a small caputegulum with a small surface texture that is arranged towards the
665:, the external nares are oriented towards the sides. The external nares are fully obscured in front view, a condition not seen in many Asian and Laramidian ankylosaurids, and are relatively small, tear-shaped openings. Positioned on the rear margin of the left nares is a small bony fragment as in
929:
and a single distal osteoderm. Each individual caudal vertebra is nearly twice as long as wide and have a spool-shaped centra. As with other ankylosaurid tail clubs, the handle is reinforced by elongated prezygapophyses which extend across for nearly half of the length of the next vertebra. The
840:
have a short but tall boot and centra that have anterior and posterior articular surfaces that are spool-shaped and are slightly amphicoelous. The neural spines of the caudal vertebrae are tall and the centra of all proximal caudal vertebra are ellipsoid in shape, with the exception of the more
977:
Only 14 noncervical osteoderms are known and are all tall with dorsal margins that are strongly keeled. Most of the osteoderms would have been positioned along the back and sides of the body, with some osteoderms possibly being positioned on the forelimbs. Of the 14 osteoderms preserved, three
815:
centra are spool-shaped and appear oblong towards the front and back as they are compressed towards the sides. However, the dorsal vertebrae of the mid-dorsal series show minor deformation. Unlike other ankylosaurids, the underside of the posterior surface of the centrum possesses a recurved,
2729:
Thomas D. Carr; Thomas E. Williamson; Brooks B. Britt; Ken
Stadtman (2011). "Evidence for high taxonomic and morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah".
2237:
Park, J. Y.; Lee, Y. N.; Currie, P. J.; Kobayashi, Y.; Koppelhus, E.; Barsbold, R.; Mateus, O.; Lee, S.; Kim, S. H. (2019). "Additional skulls of
Talarurus plicatospineus (Dinosauria: Ankylosauridae) and implications for paleobiogeography and paleoecology of armored dinosaurs".
885:
process and a partial shaft with the radial notch morphology being similar to that of other ankylosaurs. Towards the side margins, the ilium is concave and nearly straight towards the middle. Towards the middle of the ilium is a concave sulcus which forms a closed
693:
slightly taper towards the front and form a narrow, U-shaped beak. The maxillae preserves 16 alveolar cavities each. As in other ankylosaurids and nodosaurids, the maxillae are concave towards the sides and together form an hourglass configuration. Like
894:
is Y-shaped towards the sides and in medial view, and consists of a shaft that is compressed towards the sides with no anterior curvature. A small protuberance on the dorsal surface is formed between the femoral head and the greater trochanter. In
715:. The jugal forms a shelf that is ornamented towards the sides with a small ridge of rugose co-ossified bone. The quadratojugal is a small wall of bone that projects towards the front and forms the front extension of the quadratojugal horn.
903:
is very poor as it experienced severe breakage and surface weathering, it was long and narrow. The pedal phalanx is transversely wider than long. In addition, the pedal phalanx is concave along the front and back articular surfaces.
996:. Type 5 osteoderms are represented by a singular, fragmentary osteoderm that has an off-centred straight keel and a round, flat base. The overall morphology of the osteoderm resembles the osteoderms that are positioned dorsally in
2781:
706:
have an ornamented side surface that forms the rear border of the orbit and is continuous with the posterodorsal ornamentation of the jugal. The jugals consist of a broad element that touches the front-most part of the
1047:
independent from the main radiation of
Laramidian ankylosaurids. The discovery indicates a strong case for provincialism between dinosaur populations in Northern and Southern Laramidia. Furthermore, the discovery of
935:
vertebrae is the tail club knob. The knob itself displays an overall subtriangular morphology in dorsal and ventral view, and is composed of two major osteoderms and a single minor osteoderm. As in
2915:
2836:
759:. Additionally, the quadrates are relatively robust and have a strong inclination towards the front of in side view. The predentary forms the mandibular counterpart to the premaxillary
659:
has a unique suite of cranial ornamentation that consists of several symmetrical rows of small pyramidal and conical caputegulae along the dorsolateral surface of the skull. As in
947:, the major osteoderms are longer from front to back than they are wide from side to side. The major osteoderms are rounded towards the side margins and are convex, unlike in
836:. The sacral ribs have a much less pronounced T-shaped cross section as the dorsal surfaces are less developed horizontally, unlike the ribs on the dorsosacral vertebrae. The
4275:
1845:
setting that was dominated by large river channels, ponds, lakes, and wetlands. The rivers flowed across the wet alluvial to coastal plain setting and were drained into the
864:
and projects exactly upright to the side surfaces of the scapular blade. The scapular blade has a distinct surface on the ventral surface that would have supported the
555:
half rings, and dorsal and lateral osteoderms of various sizes and morphologies. It represents about 45% of the skeletal elements and is part of the collection of the
763:
and articulates with the mandibular symphysis. In addition, the rear end of the predentary has a smooth surface texture. As in other ankylosaurids, the predentary of
421:
2406:
Gates, Terry A.; Lund, Eric K.; Boyd, C. A.; DeBlieux, Donald D.; Titus, Alan L.; Evans, David C.; Getty, Michael A.; Kirkland, James I.; Eaton, Jeffrey G. (2013).
2273:
G. Frauenfelder, Timothy; R. Bell, Phil; Brougham, Tom; J. Bevitt, Joseph; D. C. Bicknell, Russell; P. Kear, Benjamin; Wroe, Stephen; E. Campione, Nicolás (2022).
432:. The Horse Mountain Gryposaur Quarry represents a multitaxic and multidominant bonebed that has produced a mostly complete, partially articulated skeleton of the
2408:"Ornithopod Dinosaurs from the Grand Staircase–Escalante National Monument Region, Utah, and Their Role in Paleobiogeographic and Macroevolutionary Studies"
2803:
Hutchison, J. Howard; Michael J. Knell; Donald B. Brinkman (2013). "Turtles from the
Kaiparowits Formation, Utah". In Alan L. Titus, Mark A. Loewen (ed.).
677:
which suggests the presence of internarial apertures. The external nares have upper and front margins that are bounded by the supranarial caputegulum. The
2923:
2782:
A preliminary report on the theropod dinosaur fauna of the late
Campanian Kaiparowits Formation, Grand Staircase-Escalante National Monument, Utah.
1043:
that is nested within Asian taxa rather than other
Laramidian taxa and suggested that this clade was a separate biogeographic dispersal event from
4158:
2566:
Taxonomic revision of latest
Cretaceous North american basal neonithischian taxa and a phylogenetic analysis of basal ornithischian relationships
2944:
2308:"The Kaiparowits Formation: A Remarkable Record of Late Cretaceous Terrestrial Environments, Ecosystems, and Evolution in Western North America"
382:, including a complete skull, tail club, a number of osteoderms, limb elements and part of its pelvis, among other remains. It was described in
4285:
3113:
590:, honours Randy Johnson, a volunteer preparator at the Natural History Museum of Utah, who prepared the skull and lower jaws of the holotype.
2898:
2812:
2139:
3012:"A large pterosaur limb bone from the Kaiparowits Formation (late Campanian) of Grand Staircase–Escalante National Monument, Utah, USA"
1833:. The Kaiparowits Formation preserves a unique record of Late Cretaceous terrestrial vertebrate ecosystems in the Western Interior of
1056:, created when dropping sea levels allowed migrations between the continents via the Beringian Land Bridge during or earlier than the
2419:
2133:
2580:
Zanno, L. E.; Sampson, S. D. (2005). "A new oviraptorosaur (Theropoda; Maniraptora) from the Late
Cretaceous (Campanian) of Utah".
877:
large yet round protuberance which forms the articulation surface for the muscle M. latissimus dorsi. The only remains of the left
784:
4175:
2435:
Sampson, S. D.; Loewen, M. A.; Farke, A. A.; Roberts, E. M.; Forster, C. A.; Smith, J. A.; Titus, A. L.; Stepanova, A. (2010).
2097:
755:
have a shaft that expands to the sides and the middle to touch the middle face of the quadratojugals and the side faces of the
2890:
556:
477:
1809:
stage, 76.26 ± 0.10 Ma. The age of the
Kaiparowits Formation makes it contemporaneous with dinosaur-bearing strata from the
816:
hook-like projection, which is followed by a longitudinal keel. Some of the dorsal vertebrae preserve portions of the fused
2916:"Crocodyliforms from the late cretaceous of grand staircase–escalante national monument and vicinity, Southern Utah, U.S.A"
4290:
4153:
1300:
3059:
135:
2275:"New Ankylosaurian Cranial Remains From the Lower Cretaceous (Upper Albian) Toolebuc Formation of Queensland, Australia"
970:
953:. The minor osteoderm is far smaller than the major osteoderms, transversely wider than it is long, and diamond-shaped.
603:
3106:
1849:. The climate was wet and humid, and supported an abundant and diverse range of organisms. The presence of aquatic
735:
472:. The age of the layer is 76.26 ± 0.10 million years based on zircon dating. In 2014, the skull was subjected to a
420:
was excavated during the 2008, 2009, and 2010 field seasons from the Horse
Mountain Gryposaur (HMG) Quarry in the
2593:
899:
and all other Late Cretaceous Laramidian ankylosaurids, the femoral head is oriented horizontally. Although the
3204:
2837:"A new species of baenid turtle from the Kaiparowits Formation (Upper Cretaceous, Campanian) of southern Utah"
930:
elongated prezygapophyses also interlock with the shorter postzygapophyses to help reinforce the handle. Each
868:
M. triceps longus caudalis. The scapular blade is paddle-shaped with a convex distal margin in side view. The
718:
2534:
2496:"A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia"
1846:
1810:
2979:(Crocodylia: Alligatoroidea) from the Upper Cretaceous Kaiparowits Formation (late Campanian) of Utah, USA"
1791:
4202:
2549:
The evolution and biogeography of ankylosaurid dinosaurs from the late cretaceous of western north america
2120:
Loewen, Mark A.; Burns, Michael E.; Getty, Michael A.; Kirkland, James I.; Vickaryous, Matthew K. (2013).
1818:
1814:
974:
half rings are bulbous and oval, and accommodate the secondary osteoderms, which were also oval in shape.
4247:
3231:
3099:
1365:
698:, the lacrimals are ornamented with a tetrahedral-shaped lacrimal caputegulum, although the lacrimal in
469:
405:
369:
311:
586:" (head), in reference to the thorn-like cranial caputegulae of the holotype skull. The specific name,
386:
by Jelle P. Wiersma and Randall B. Irmis. It is closely related and shares similar cranial anatomy to
4280:
4234:
4185:
2952:
2848:
2739:
2680:"The most complete enantiornithine from North America and a phylogenetic analysis of the Avisauridae"
2632:
2448:
2092:
1853:
and abundance of aquatic vertebrates and plants testifies to the wet nature of the alluvial setting.
383:
288:
2615:
Lindsay E. Zanno; David J. Varricchio; Patrick M. O'Connor; Alan L. Titus; Michael J. Knell (2011).
3591:
3488:
2306:
M. Roberts, Eric; D. Sampson, Scott; L. Deino, Alan; A. Bowring, Samuel; Buchwaldt, Robert (2013).
1822:
2122:"Review of Late Cretaceous Ankylosaurian Dinosaurs from the Grand Staircase Region, Southern Utah"
2057:
611:
3327:
2887:
Non-dinosaurian lower vertebrates across the Cretaceous-Tertiary boundary in northeastern Montana
2864:
2763:
2597:
2385:
2354:
2255:
2044:
1882:
1826:
812:
804:
708:
130:
4252:
2019:
4084:
3348:
3043:
2894:
2808:
2755:
2711:
2660:
2525:
2476:
2415:
2219:
2129:
1444:
760:
661:
632:
425:
388:
1995:
1805:
is known from the middle unit of the Kaiparowits Formation which has been dated to the upper
3549:
3496:
3334:
3073:
3033:
3023:
2990:
2856:
2747:
2701:
2691:
2650:
2640:
2589:
2515:
2507:
2466:
2456:
2346:
2286:
2247:
2209:
2199:
1937:
1511:
1413:
703:
643:
4014:
3978:
3918:
3884:
3877:
3770:
3711:
3583:
3480:
3396:
3356:
3189:
1971:
1877:
1830:
1494:
1341:
1204:
865:
768:
678:
673:
507:, dorsosacral vertebrae, sacral vertebrae, caudosacral vertebra, caudal vertebrae, dorsal
361:
30:
2852:
2743:
2636:
2564:
2452:
2437:"New horned dinosaurs from Utah provide evidence for intracontinental dinosaur endemism"
117:
4077:
4063:
3957:
3930:
3826:
3807:
3722:
3668:
3533:
3503:
3473:
3436:
3412:
3038:
3011:
2973:
Farke, Andrew A.; Henn, Madison M.; Woodward, Samuel J.; Xu, Heendong A. (2014-01-30).
2706:
2679:
2655:
2616:
2547:
2520:
2495:
2471:
2436:
2214:
2188:
gen. et sp. nov., from the upper Campanian Kaiparowits Formation of southern Utah, USA"
2183:
2062:
2024:
1980:
1959:
1954:
1941:
1921:
1917:
1914:
1909:
1904:
1886:
1872:
1838:
1638:
1570:
1276:
1166:
1146:
1140:
949:
937:
756:
628:
620:
560:
465:
417:
350:
254:
241:
795:
4269:
4040:
3950:
3943:
3895:
3839:
3786:
3752:
3653:
3599:
3574:
3556:
3459:
3405:
3378:
3370:
2601:
2407:
2358:
2337:(Dinosauria: Hadrosauridae) from the Upper Campanian Kaiparowits Formation of Utah".
2307:
2259:
1975:
1899:
1863:
1860:
1842:
1834:
1097:
1053:
861:
820:. The ribs that are fused to the dorsal vertebrae are T-shaped in cross section. The
752:
686:
575:
267:
68:
2868:
2767:
2621:
gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America"
961:
844:
787:. The prearticular comprises the retroarticular process, along with the surangular.
476:
in order to reveal the internal anatomy. The specimen was subsequently described in
378:, is known from the most complete ankylosaur specimen ever discovered from southern
4070:
4056:
3985:
3869:
3793:
3738:
3689:
3661:
3646:
3633:
3606:
3525:
3510:
3466:
3446:
3313:
3295:
3273:
3243:
3216:
3164:
3122:
2350:
2077:
2072:
2053:
2001:
1950:
1926:
1893:
1889:
1672:
1655:
1159:
A phylogenetic analysis conducted by Wiersma and Irmis (2018) is reproduced below.
1152:
1128:
1008:
943:
921:
907:
833:
667:
624:
548:
532:
454:
449:
343:
228:
196:
2860:
2251:
619:
The describing authors indicated several distinguishing traits. Some of these are
4225:
2645:
2494:
Sampson, S. D.; Lund, E. K.; Loewen, M. A.; Farke, A. A.; Clayton, K. E. (2013).
2461:
4105:
4022:
3971:
3911:
3862:
3704:
3682:
3614:
3563:
3363:
3341:
3173:
3085:
2780:
Zanno, L.E., Weirsma, J.P., Loewen, M.A., Sampson, S.D. and Getty, M.A. (2006).
2066:
2013:
2007:
1962:
1934:
1930:
1867:
1604:
1252:
1228:
1134:
1110:
992:
931:
919:
is made up of the handle, which consists of eleven fused caudal vertebrae as in
828:
and a single caudosacral vertebra, which are all fused along the centra and the
437:
433:
400:
212:
43:
856:
are curved outward towards the sides and curve inwards towards the middle. The
4098:
4091:
4029:
3904:
3855:
3847:
3800:
3745:
3731:
3697:
3675:
3541:
3385:
3320:
2889:. Volume 134 of University of California publications in geological sciences.
2751:
2291:
2274:
1587:
1317:
1180:
1089:
1084:
1000:. The surface texture of the osteoderm is smooth with small, shallow pitting.
887:
772:
690:
682:
500:
88:
53:
4112:
3992:
3779:
3759:
3621:
3518:
3420:
2974:
2080:
2048:
2029:
1945:
1806:
1389:
1072:
1066:
1057:
1036:
986:
926:
916:
882:
821:
552:
512:
462:
379:
365:
147:
93:
37:
3047:
2759:
2715:
2664:
2529:
2511:
2480:
2223:
631:
formed by the basioccipital is located obliquely above and in front of the
4219:
4119:
3155:
3146:
2034:
1988:
1850:
1621:
869:
857:
837:
829:
808:
780:
776:
520:
504:
496:
484:
458:
413:
353:
180:
167:
83:
78:
63:
58:
48:
3028:
2995:
4007:
3999:
2920:
At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah
2805:
At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah
2696:
2412:
At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah
2389:
2312:
At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah
2204:
1991:
1966:
1528:
1078:
980:
891:
873:
853:
712:
524:
516:
473:
98:
73:
2126:
At the Top of the Grand Staircase-The Late Cretaceous of Southern Utah
1064:(2019) found both Akainacephalus and Nodocephalosaurus to be basal to
702:
is well defined in lateral view and has a rugose surface texture. The
368:, 76.26 Ma) in what is now the Horse Mountain Gryposaur Quarry of the
3140:
2039:
1798:(lower left) with contemporary dinosaurs of the Kaiparowits Formation
900:
825:
544:
445:
442:
157:
4196:
2121:
1108:
as sister taxon to two clades, with one of the clades consisting of
2678:
Jessie Atterholt; J. Howard Hutchison; Jingmai K. O’Connor (2018).
468:
that is located within the lower section of the middle unit of the
3964:
1790:
1007:
960:
906:
843:
794:
734:
717:
642:
610:
602:
567:
540:
536:
492:
399:
346:
4239:
4126:
1093:
1044:
878:
528:
429:
357:
4200:
3133:
3095:
3091:
1052:
also indicates at least two faunal migrations between Asia and
817:
508:
1016:(left) compared to its close relatives (from left to right):
1122:
while the other consists of more deeply nested taxa such as
1092:
stage and that ankylosaurines dispersed at least twice from
2594:
10.1671/0272-4634(2005)025[0897:anotmf]2.0.co;2
2786:
Learning from the Land Symposium: Geology and Paleontology
872:
is square in shape and is fused to the right scapula. The
2945:"First Report of the Hyper-Giant Cretaceous Crocodylian
461:. The bonebed is deposited in a fine- to medium-grained
404:
Map of the Grand Staircase–Escalante National Monument.
2500:
Proceedings of the Royal Society B: Biological Sciences
2333:
Gates, Terry; Sampson, Scott (2006). "A new species of
1088:, which suggests that a migration occurred before the
3057:
2885:
Bryant, Laurie J. (1989). "Systematic Paleontology".
965:
Cervical half rings and other noncervical osteoderms.
767:
is transversely almost straight. The total number of
16:
Ankylosaurid dinosaur genus from Late Cretaceous Utah
4209:
4039:
3929:
3894:
3825:
3769:
3721:
3632:
3573:
3445:
3434:
3395:
3293:
3242:
3215:
3188:
925:, and the knob, which consists of only two lateral
3010:A. Farke, Andrew (2021). Piñeiro, Graciela (ed.).
2798:
2796:
2794:
860:of the scapula is positioned directly above the
615:Preserved elements and skeletal reconstructions
2807:. Indiana University Press. pp. 295–318.
2414:. Indiana University Press. pp. 463–481.
2177:
2175:
2173:
2171:
2169:
2128:. Indiana University Press. pp. 445–462.
771:is unknown as they have been eroded away. The
3107:
2880:
2878:
2410:. In Titus, Alan L.; Loewen, Mark A. (eds.).
2310:. In L. Titus, Alan; A. Loewen, Mark (eds.).
2167:
2165:
2163:
2161:
2159:
2157:
2155:
2153:
2151:
2149:
2124:. In Titus, Alan L.; Loewen, Mark A. (eds.).
811:and a distinct keel on the ventral side. The
8:
2788:. Washington, DC: Bureau of Land Management.
2569:(Thesis). The University of Texas at Austin.
2115:
2113:
1859:coexisted with other dinosaurs, such as the
779:, contributes to the tallest portion of the
2401:
2399:
2182:Jelle P. Wiersma; Randall B. Irmis (2018).
422:Grand Staircase–Escalante National Monument
4276:Late Cretaceous dinosaurs of North America
4197:
3935:
3831:
3638:
3451:
3442:
3305:
3301:
3290:
3221:
3194:
3185:
3130:
3114:
3100:
3092:
480:by Jelle P. Wiersma and Randall B. Irmis.
116:
20:
3037:
3027:
2994:
2830:
2828:
2826:
2824:
2705:
2695:
2654:
2644:
2519:
2470:
2460:
2290:
2213:
2203:
2184:"A new southern Laramidian ankylosaurid,
1983:. Non-dinosaur taxa contemporaneous with
890:. As with other ankylosaurids, the right
775:, along with the posterior margin of the
2372:Weishampel, D.B.; Jensen, J.A. (1979). "
689:due to anteroposterior deformation. The
3064:
2109:
2376:(Reptilia: Hadrosauridae) from Utah".
848:Tibia in anterior and posterior views.
453:, an articulated skeleton of a small
7:
824:consists of four dorsosacral, three
1562:
1486:
1436:
1429:
1405:
1381:
1357:
1333:
1292:
1268:
1244:
1220:
1196:
1172:
1162:
2914:Irmis, Randall B. (January 2013).
2841:Journal of Vertebrate Paleontology
2582:Journal of Vertebrate Paleontology
2339:Journal of Vertebrate Paleontology
14:
799:Sacrum and caudosacral vertebrae.
457:, and a partial skull of a small
4181:
4180:
4157:
4152:
3079:
3067:
2142:from the original on 2022-07-12.
134:
2098:Timeline of ankylosaur research
1031:Wiersma and Irmis (2018) found
2891:University of California Press
2546:Wiersma, Jelle Pouwel (2016).
2351:10.1080/02724634.2006.10010069
1878:Parasaurolophus cyrtocristatus
557:Natural History Museum of Utah
1:
4286:Fossil taxa described in 2018
2861:10.1080/02724634.2015.1009084
2552:(Thesis). University of Utah.
2252:10.1016/j.cretres.2019.104340
1837:as it was deposited in a wet
1301:Pinacosaurus mephistocephalus
372:. The type and only species,
2646:10.1371/journal.pone.0024487
2617:"A new troodontid theropod,
2462:10.1371/journal.pone.0012292
342:(meaning "thorn head") is a
2563:Boyd, Clint Aaroen (2012).
1012:The skull ornamentation of
711:and the rear margin of the
681:are almond-shaped and both
4307:
2835:Lively, Joshua R. (2015).
2279:Frontiers in Earth Science
911:Tail club handle and knob.
4170:
4150:
3938:
3834:
3641:
3454:
3308:
3304:
3289:
3270:
3224:
3197:
3184:
3129:
2922:: 424–444. Archived from
2752:10.1007/s00114-011-0762-7
2292:10.3389/feart.2022.803505
1868:Gryposaurus monumentensis
1669:
1652:
1635:
1618:
1601:
1584:
1567:
1560:
1525:
1508:
1491:
1484:
1458:
1441:
1434:
1427:
1410:
1403:
1386:
1379:
1362:
1355:
1338:
1331:
1314:
1297:
1290:
1273:
1266:
1249:
1242:
1225:
1218:
1201:
1194:
1177:
1170:
722:Mandibles and predentary.
317:
310:
131:Scientific classification
129:
124:
115:
23:
1035:to be within a southern
582:" (thorn or spine) and "
2378:Journal of Paleontology
2186:Akainacephalus johnsoni
2076:, and an indeterminate
1979:, and an indeterminate
1847:Western Interior Seaway
685:are slightly above the
375:Akainacephalus johnsoni
330:Wiersma and Irmis, 2018
324:Akainacephalus johnsoni
125:Reconstructed skeleton
2512:10.1098/rspb.2013.1186
1799:
1028:
966:
912:
849:
800:
748:
723:
652:
616:
608:
574:, is derived from the
441:, the holotype of the
409:
360:that lived during the
4248:Paleobiology Database
1794:
1366:Pinacosaurus grangeri
1060:stage. However, Park
1011:
964:
910:
847:
798:
738:
721:
646:
614:
606:
599:Distinguishing traits
470:Kaiparowits Formation
406:Kaiparowits Formation
403:
370:Kaiparowits Formation
4291:Ornithischian genera
2093:2018 in paleontology
1940:"Skaladromeus", the
396:Discovery and naming
3029:10.7717/peerj.10766
2996:10.5070/P9303016247
2853:2015JVPal..35E9084L
2744:2011NW.....98..241C
2732:Naturwissenschaften
2637:2011PLoSO...624487Z
2453:2010PLoSO...512292S
2240:Cretaceous Research
1929:, an indeterminate
1925:, an indeterminate
1881:, an indeterminate
807:has a spool-shaped
607:Life reconstruction
551:, partial cervical
412:An almost complete
287:Wiersma and Irmis,
2893:. pp. 10–58.
2697:10.7717/peerj.5910
2506:(1766): 20131186.
2205:10.7717/peerj.5016
1800:
1029:
971:cervical half ring
969:Both the anterior
967:
913:
850:
801:
749:
724:
653:
617:
609:
410:
4263:
4262:
4203:Taxon identifiers
4194:
4193:
4166:
4165:
4148:
4147:
4144:
4143:
4140:
4139:
4136:
4135:
4085:Nodocephalosaurus
3821:
3820:
3817:
3816:
3430:
3429:
3349:Serendipaceratops
3285:
3284:
3281:
3280:
3266:
3265:
3262:
3261:
2900:978-0-520-09735-3
2814:978-0-253-00896-1
2345:(3, Suppl): 65A.
2314:. pp. 85–106
2054:soft shell turtle
1883:brachylophosaurin
1783:
1782:
1774:
1773:
1765:
1764:
1756:
1755:
1747:
1746:
1738:
1737:
1729:
1728:
1720:
1719:
1711:
1710:
1702:
1701:
1693:
1692:
1684:
1683:
1549:
1548:
1540:
1539:
1473:
1472:
1445:Nodocephalosaurus
1116:Nodocephalosaurus
1104:(2022) recovered
1041:Nodocephalosaurus
1039:clade containing
1018:Nodocephalosaurus
805:cervical vertebra
745:Nodocephalosaurus
700:Nodocephalosaurus
696:Nodocephalosaurus
662:Nodocephalosaurus
633:occipital condyle
408:is in dark green.
389:Nodocephalosaurus
335:
334:
291:
4298:
4256:
4255:
4243:
4242:
4230:
4229:
4228:
4198:
4184:
4183:
4161:
4156:
3936:
3832:
3639:
3550:Propanoplosaurus
3497:Glyptodontopelta
3452:
3443:
3335:Priodontognathus
3306:
3302:
3291:
3222:
3195:
3186:
3179:
3178:
3131:
3116:
3109:
3102:
3093:
3084:
3083:
3082:
3072:
3071:
3070:
3063:
3052:
3051:
3041:
3031:
3007:
3001:
3000:
2998:
2970:
2964:
2963:
2961:
2960:
2951:. Archived from
2941:
2935:
2934:
2932:
2931:
2911:
2905:
2904:
2882:
2873:
2872:
2832:
2819:
2818:
2800:
2789:
2778:
2772:
2771:
2726:
2720:
2719:
2709:
2699:
2675:
2669:
2668:
2658:
2648:
2612:
2606:
2605:
2577:
2571:
2570:
2560:
2554:
2553:
2543:
2537:
2533:
2523:
2491:
2485:
2484:
2474:
2464:
2432:
2426:
2425:
2403:
2394:
2393:
2384:(6): 1422–1427.
2369:
2363:
2362:
2330:
2324:
2323:
2321:
2319:
2303:
2297:
2296:
2294:
2270:
2264:
2263:
2234:
2228:
2227:
2217:
2207:
2179:
2144:
2143:
2117:
2045:nanhsiungchelyid
1787:Paleoenvironment
1563:
1512:Tarchia kielanae
1487:
1437:
1430:
1414:Tarchia gigantea
1406:
1382:
1358:
1334:
1293:
1269:
1245:
1221:
1197:
1173:
1163:
858:acromion process
838:caudal vertebrae
826:sacral vertebrae
813:dorsal vertebrae
785:coronoid process
491:, consists of a
326:
322:
303:A. johnsoni
286:
279:
266:
253:
240:
227:
211:
195:
139:
138:
120:
110:
40:
29:Temporal range:
21:
4306:
4305:
4301:
4300:
4299:
4297:
4296:
4295:
4266:
4265:
4264:
4259:
4251:
4246:
4238:
4233:
4224:
4223:
4218:
4205:
4195:
4190:
4162:
4132:
4035:
4015:Tianzhenosaurus
3979:Minotaurasaurus
3925:
3919:Zhongyuansaurus
3890:
3885:Zhejiangosaurus
3878:Liaoningosaurus
3813:
3771:Struthiosaurini
3765:
3717:
3712:Tatankacephalus
3628:
3584:Gargoyleosaurus
3569:
3481:Gargoyleosaurus
3439:
3426:
3397:Parankylosauria
3391:
3357:Sinankylosaurus
3298:
3277:
3258:
3238:
3211:
3205:Avemetatarsalia
3190:Avemetatarsalia
3180:
3136:
3135:
3125:
3120:
3090:
3080:
3078:
3068:
3066:
3058:
3056:
3055:
3009:
3008:
3004:
2972:
2971:
2967:
2958:
2956:
2943:
2942:
2938:
2929:
2927:
2913:
2912:
2908:
2901:
2884:
2883:
2876:
2847:(6): e1009084.
2834:
2833:
2822:
2815:
2802:
2801:
2792:
2779:
2775:
2728:
2727:
2723:
2677:
2676:
2672:
2614:
2613:
2609:
2579:
2578:
2574:
2562:
2561:
2557:
2545:
2544:
2540:
2493:
2492:
2488:
2434:
2433:
2429:
2422:
2405:
2404:
2397:
2374:Parasaurolophus
2371:
2370:
2366:
2332:
2331:
2327:
2317:
2315:
2305:
2304:
2300:
2272:
2271:
2267:
2236:
2235:
2231:
2181:
2180:
2147:
2136:
2119:
2118:
2111:
2106:
2089:
1960:enantiornithine
1831:Aguja Formation
1789:
1784:
1775:
1766:
1757:
1748:
1739:
1730:
1721:
1712:
1703:
1694:
1685:
1550:
1541:
1495:Minotaurasaurus
1474:
1342:Tianzhenosaurus
1205:Zhongyuansaurus
1100:. Frauenfelder
1026:Minotaurasaurus
1006:
959:
793:
674:Minotaurasaurus
641:
601:
596:
398:
362:Late Cretaceous
331:
328:
320:
319:
306:
285:
277:
264:
251:
238:
225:
209:
193:
133:
111:
109:
108:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
35:
34:
31:Late Cretaceous
27:
17:
12:
11:
5:
4304:
4302:
4294:
4293:
4288:
4283:
4278:
4268:
4267:
4261:
4260:
4258:
4257:
4244:
4231:
4215:
4213:
4211:Akainacephalus
4207:
4206:
4201:
4192:
4191:
4189:
4188:
4178:
4171:
4168:
4167:
4164:
4163:
4151:
4149:
4146:
4145:
4142:
4141:
4138:
4137:
4134:
4133:
4131:
4130:
4123:
4116:
4109:
4102:
4095:
4088:
4081:
4078:Euoplocephalus
4074:
4067:
4064:Anodontosaurus
4060:
4053:
4050:Akainacephalus
4045:
4043:
4037:
4036:
4034:
4033:
4026:
4019:
4011:
4004:
3996:
3989:
3982:
3975:
3968:
3961:
3958:Crichtonsaurus
3954:
3947:
3939:
3933:
3931:Ankylosaurinae
3927:
3926:
3924:
3923:
3915:
3908:
3900:
3898:
3892:
3891:
3889:
3888:
3881:
3874:
3866:
3859:
3852:
3844:
3835:
3829:
3827:Ankylosauridae
3823:
3822:
3819:
3818:
3815:
3814:
3812:
3811:
3808:Struthiosaurus
3804:
3797:
3790:
3783:
3775:
3773:
3767:
3766:
3764:
3763:
3756:
3749:
3742:
3735:
3727:
3725:
3723:Panoplosaurini
3719:
3718:
3716:
3715:
3708:
3701:
3694:
3686:
3679:
3672:
3669:Niobrarasaurus
3665:
3658:
3650:
3642:
3636:
3630:
3629:
3627:
3626:
3618:
3611:
3603:
3596:
3588:
3579:
3577:
3571:
3570:
3568:
3567:
3560:
3553:
3546:
3538:
3534:Polacanthoides
3530:
3522:
3515:
3507:
3504:Horshamosaurus
3500:
3493:
3485:
3477:
3474:Dongyangopelta
3470:
3463:
3455:
3449:
3440:
3437:Euankylosauria
3435:
3432:
3431:
3428:
3427:
3425:
3424:
3417:
3413:Kunbarrasaurus
3409:
3401:
3399:
3393:
3392:
3390:
3389:
3382:
3375:
3367:
3360:
3353:
3345:
3338:
3331:
3324:
3317:
3309:
3299:
3294:
3287:
3286:
3283:
3282:
3279:
3278:
3271:
3268:
3267:
3264:
3263:
3260:
3259:
3257:
3256:
3255:
3254:
3248:
3246:
3240:
3239:
3237:
3236:
3235:
3234:
3225:
3219:
3213:
3212:
3210:
3209:
3208:
3207:
3198:
3192:
3182:
3181:
3177:
3176:
3167:
3158:
3149:
3143:
3134:
3127:
3126:
3121:
3119:
3118:
3111:
3104:
3096:
3089:
3088:
3076:
3054:
3053:
3002:
2965:
2936:
2906:
2899:
2874:
2820:
2813:
2790:
2773:
2738:(3): 241–246.
2721:
2670:
2619:Talos sampsoni
2607:
2588:(4): 897–904.
2572:
2555:
2538:
2486:
2427:
2420:
2395:
2364:
2325:
2298:
2265:
2229:
2145:
2134:
2108:
2107:
2105:
2102:
2101:
2100:
2095:
2088:
2085:
2025:paracryptodire
1985:Akainacephalus
1938:thescelosaurid
1922:Anodontosaurus
1910:Nasutoceratops
1857:Akainacephalus
1803:Akainacephalus
1796:Akainacephalus
1788:
1785:
1781:
1780:
1777:
1776:
1772:
1771:
1768:
1767:
1763:
1762:
1759:
1758:
1754:
1753:
1750:
1749:
1745:
1744:
1741:
1740:
1736:
1735:
1732:
1731:
1727:
1726:
1723:
1722:
1718:
1717:
1714:
1713:
1709:
1708:
1705:
1704:
1700:
1699:
1696:
1695:
1691:
1690:
1687:
1686:
1682:
1681:
1678:
1677:
1668:
1665:
1664:
1661:
1660:
1651:
1648:
1647:
1644:
1643:
1639:Anodontosaurus
1634:
1631:
1630:
1627:
1626:
1617:
1614:
1613:
1610:
1609:
1600:
1597:
1596:
1593:
1592:
1583:
1580:
1579:
1576:
1575:
1571:Euoplocephalus
1566:
1561:
1559:
1556:
1555:
1552:
1551:
1547:
1546:
1543:
1542:
1538:
1537:
1534:
1533:
1524:
1521:
1520:
1517:
1516:
1507:
1504:
1503:
1500:
1499:
1490:
1485:
1483:
1480:
1479:
1476:
1475:
1471:
1470:
1467:
1466:
1462:Akainacephalus
1457:
1454:
1453:
1450:
1449:
1440:
1435:
1433:
1428:
1426:
1423:
1422:
1419:
1418:
1409:
1404:
1402:
1399:
1398:
1395:
1394:
1385:
1380:
1378:
1375:
1374:
1371:
1370:
1361:
1356:
1354:
1351:
1350:
1347:
1346:
1337:
1332:
1330:
1327:
1326:
1323:
1322:
1313:
1310:
1309:
1306:
1305:
1296:
1291:
1289:
1286:
1285:
1282:
1281:
1277:Crichtonsaurus
1272:
1267:
1265:
1262:
1261:
1258:
1257:
1248:
1243:
1241:
1238:
1237:
1234:
1233:
1224:
1219:
1217:
1214:
1213:
1210:
1209:
1200:
1195:
1193:
1190:
1189:
1186:
1185:
1176:
1171:
1169:
1167:Ankylosauridae
1161:
1147:Euoplocephalus
1141:Anodontosaurus
1106:Akainacephalus
1050:Akainacephalus
1033:Akainacephalus
1014:Akainacephalus
1005:
1004:Classification
1002:
958:
955:
950:Anodontosaurus
938:Euoplocephalus
897:Akainacephalus
803:The posterior
792:
789:
783:and forms the
765:Akainacephalus
741:Akainacephalus
739:Comparison of
729:Akainacephalus
657:Akainacephalus
651:in side views.
649:Akainacephalus
640:
637:
629:foramen magnum
621:autapomorphies
600:
597:
595:
592:
572:Akainacephalus
561:Salt Lake City
466:crevasse splay
397:
394:
356:from southern
339:Akainacephalus
333:
332:
329:
315:
314:
308:
307:
299:
297:
293:
292:
282:Akainacephalus
275:
271:
270:
262:
258:
257:
255:Ankylosaurinae
249:
245:
244:
242:Ankylosauridae
236:
232:
231:
223:
216:
215:
207:
200:
199:
191:
184:
183:
178:
171:
170:
165:
161:
160:
155:
151:
150:
145:
141:
140:
127:
126:
122:
121:
113:
112:
104:
103:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
41:
28:
25:Akainacephalus
15:
13:
10:
9:
6:
4:
3:
2:
4303:
4292:
4289:
4287:
4284:
4282:
4279:
4277:
4274:
4273:
4271:
4254:
4249:
4245:
4241:
4236:
4232:
4227:
4221:
4217:
4216:
4214:
4212:
4208:
4204:
4199:
4187:
4179:
4177:
4173:
4172:
4169:
4160:
4155:
4129:
4128:
4124:
4122:
4121:
4117:
4115:
4114:
4110:
4108:
4107:
4103:
4101:
4100:
4096:
4094:
4093:
4089:
4087:
4086:
4082:
4080:
4079:
4075:
4073:
4072:
4068:
4066:
4065:
4061:
4059:
4058:
4054:
4052:
4051:
4047:
4046:
4044:
4042:
4041:Ankylosaurini
4038:
4032:
4031:
4027:
4025:
4024:
4020:
4017:
4016:
4012:
4010:
4009:
4005:
4002:
4001:
3997:
3995:
3994:
3990:
3988:
3987:
3983:
3981:
3980:
3976:
3974:
3973:
3969:
3967:
3966:
3962:
3960:
3959:
3955:
3953:
3952:
3951:Crichtonpelta
3948:
3946:
3945:
3944:Bissektipelta
3941:
3940:
3937:
3934:
3932:
3928:
3921:
3920:
3916:
3914:
3913:
3909:
3907:
3906:
3902:
3901:
3899:
3897:
3896:Shamosaurinae
3893:
3887:
3886:
3882:
3880:
3879:
3875:
3872:
3871:
3867:
3865:
3864:
3860:
3858:
3857:
3853:
3850:
3849:
3845:
3842:
3841:
3840:Ahshislepelta
3837:
3836:
3833:
3830:
3828:
3824:
3810:
3809:
3805:
3803:
3802:
3798:
3796:
3795:
3791:
3789:
3788:
3787:Hungarosaurus
3784:
3782:
3781:
3777:
3776:
3774:
3772:
3768:
3762:
3761:
3757:
3755:
3754:
3753:Panoplosaurus
3750:
3748:
3747:
3743:
3741:
3740:
3736:
3734:
3733:
3729:
3728:
3726:
3724:
3720:
3714:
3713:
3709:
3707:
3706:
3702:
3700:
3699:
3695:
3692:
3691:
3687:
3685:
3684:
3680:
3678:
3677:
3673:
3671:
3670:
3666:
3664:
3663:
3659:
3656:
3655:
3654:Ahshislepelta
3651:
3649:
3648:
3644:
3643:
3640:
3637:
3635:
3631:
3624:
3623:
3619:
3617:
3616:
3612:
3609:
3608:
3604:
3602:
3601:
3600:Hoplitosaurus
3597:
3594:
3593:
3589:
3586:
3585:
3581:
3580:
3578:
3576:
3575:Polacanthinae
3572:
3566:
3565:
3561:
3559:
3558:
3557:Rhadinosaurus
3554:
3552:
3551:
3547:
3544:
3543:
3539:
3536:
3535:
3531:
3528:
3527:
3523:
3521:
3520:
3516:
3513:
3512:
3508:
3506:
3505:
3501:
3499:
3498:
3494:
3491:
3490:
3486:
3483:
3482:
3478:
3476:
3475:
3471:
3469:
3468:
3464:
3462:
3461:
3460:Acanthopholis
3457:
3456:
3453:
3450:
3448:
3444:
3441:
3438:
3433:
3423:
3422:
3418:
3415:
3414:
3410:
3408:
3407:
3406:Antarctopelta
3403:
3402:
3400:
3398:
3394:
3388:
3387:
3383:
3381:
3380:
3379:Tianchisaurus
3376:
3373:
3372:
3371:Stegosaurides
3368:
3366:
3365:
3361:
3359:
3358:
3354:
3351:
3350:
3346:
3344:
3343:
3339:
3337:
3336:
3332:
3330:
3329:
3325:
3323:
3322:
3318:
3316:
3315:
3311:
3310:
3307:
3303:
3300:
3297:
3292:
3288:
3276:
3275:
3269:
3252:
3251:
3250:
3249:
3247:
3245:
3241:
3233:
3229:
3228:
3227:
3226:
3223:
3220:
3218:
3214:
3206:
3202:
3201:
3200:
3199:
3196:
3193:
3191:
3187:
3183:
3175:
3171:
3168:
3166:
3162:
3159:
3157:
3153:
3150:
3148:
3144:
3142:
3138:
3137:
3132:
3128:
3124:
3117:
3112:
3110:
3105:
3103:
3098:
3097:
3094:
3087:
3077:
3075:
3065:
3061:
3049:
3045:
3040:
3035:
3030:
3025:
3021:
3017:
3013:
3006:
3003:
2997:
2992:
2988:
2984:
2980:
2978:
2969:
2966:
2955:on 2017-07-29
2954:
2950:
2948:
2940:
2937:
2926:on 2017-11-07
2925:
2921:
2917:
2910:
2907:
2902:
2896:
2892:
2888:
2881:
2879:
2875:
2870:
2866:
2862:
2858:
2854:
2850:
2846:
2842:
2838:
2831:
2829:
2827:
2825:
2821:
2816:
2810:
2806:
2799:
2797:
2795:
2791:
2787:
2783:
2777:
2774:
2769:
2765:
2761:
2757:
2753:
2749:
2745:
2741:
2737:
2733:
2725:
2722:
2717:
2713:
2708:
2703:
2698:
2693:
2689:
2685:
2681:
2674:
2671:
2666:
2662:
2657:
2652:
2647:
2642:
2638:
2634:
2631:(9): e24487.
2630:
2626:
2622:
2620:
2611:
2608:
2603:
2599:
2595:
2591:
2587:
2583:
2576:
2573:
2568:
2567:
2559:
2556:
2551:
2550:
2542:
2539:
2536:
2531:
2527:
2522:
2517:
2513:
2509:
2505:
2501:
2497:
2490:
2487:
2482:
2478:
2473:
2468:
2463:
2458:
2454:
2450:
2447:(9): e12292.
2446:
2442:
2438:
2431:
2428:
2423:
2421:9780253008961
2417:
2413:
2409:
2402:
2400:
2396:
2391:
2387:
2383:
2379:
2375:
2368:
2365:
2360:
2356:
2352:
2348:
2344:
2340:
2336:
2329:
2326:
2313:
2309:
2302:
2299:
2293:
2288:
2284:
2280:
2276:
2269:
2266:
2261:
2257:
2253:
2249:
2245:
2241:
2233:
2230:
2225:
2221:
2216:
2211:
2206:
2201:
2197:
2193:
2189:
2187:
2178:
2176:
2174:
2172:
2170:
2168:
2166:
2164:
2162:
2160:
2158:
2156:
2154:
2152:
2150:
2146:
2141:
2137:
2135:9780253008961
2131:
2127:
2123:
2116:
2114:
2110:
2103:
2099:
2096:
2094:
2091:
2090:
2086:
2084:
2082:
2079:
2075:
2074:
2069:
2068:
2064:
2063:alligatoroids
2060:
2059:
2055:
2051:
2050:
2046:
2042:
2041:
2036:
2032:
2031:
2026:
2022:
2021:
2016:
2015:
2010:
2009:
2004:
2003:
1998:
1997:
1993:
1990:
1986:
1982:
1978:
1977:
1976:Teratophoneus
1973:
1972:tyrannosaurid
1969:
1968:
1964:
1961:
1957:
1956:
1952:
1948:
1947:
1943:
1939:
1936:
1932:
1928:
1924:
1923:
1919:
1916:
1915:ankylosaurine
1912:
1911:
1906:
1905:centrosaurine
1902:
1901:
1900:Kosmoceratops
1896:
1895:
1891:
1888:
1887:chasmosaurine
1884:
1880:
1879:
1874:
1873:lambeosaurine
1870:
1869:
1865:
1862:
1858:
1854:
1852:
1848:
1844:
1843:coastal plain
1840:
1836:
1835:North America
1832:
1828:
1824:
1820:
1816:
1812:
1811:Dinosaur Park
1808:
1804:
1797:
1793:
1786:
1779:
1778:
1770:
1769:
1761:
1760:
1752:
1751:
1743:
1742:
1734:
1733:
1725:
1724:
1716:
1715:
1707:
1706:
1698:
1697:
1689:
1688:
1680:
1679:
1676:
1675:
1674:
1667:
1666:
1663:
1662:
1659:
1658:
1657:
1650:
1649:
1646:
1645:
1642:
1641:
1640:
1633:
1632:
1629:
1628:
1625:
1624:
1623:
1616:
1615:
1612:
1611:
1608:
1607:
1606:
1599:
1598:
1595:
1594:
1591:
1590:
1589:
1582:
1581:
1578:
1577:
1574:
1573:
1572:
1565:
1564:
1558:
1557:
1554:
1553:
1545:
1544:
1536:
1535:
1532:
1531:
1530:
1523:
1522:
1519:
1518:
1515:
1514:
1513:
1506:
1505:
1502:
1501:
1498:
1497:
1496:
1489:
1488:
1482:
1481:
1478:
1477:
1469:
1468:
1465:
1464:
1463:
1456:
1455:
1452:
1451:
1448:
1447:
1446:
1439:
1438:
1432:
1431:
1425:
1424:
1421:
1420:
1417:
1416:
1415:
1408:
1407:
1401:
1400:
1397:
1396:
1393:
1392:
1391:
1384:
1383:
1377:
1376:
1373:
1372:
1369:
1368:
1367:
1360:
1359:
1353:
1352:
1349:
1348:
1345:
1344:
1343:
1336:
1335:
1329:
1328:
1325:
1324:
1321:
1320:
1319:
1312:
1311:
1308:
1307:
1304:
1303:
1302:
1295:
1294:
1288:
1287:
1284:
1283:
1280:
1279:
1278:
1271:
1270:
1264:
1263:
1260:
1259:
1256:
1255:
1254:
1247:
1246:
1240:
1239:
1236:
1235:
1232:
1231:
1230:
1223:
1222:
1216:
1215:
1212:
1211:
1208:
1207:
1206:
1199:
1198:
1192:
1191:
1188:
1187:
1184:
1183:
1182:
1175:
1174:
1168:
1165:
1164:
1160:
1157:
1155:
1154:
1149:
1148:
1143:
1142:
1137:
1136:
1131:
1130:
1125:
1121:
1117:
1113:
1112:
1107:
1103:
1099:
1098:North America
1095:
1091:
1087:
1086:
1081:
1080:
1075:
1074:
1069:
1068:
1063:
1059:
1055:
1054:North America
1051:
1046:
1042:
1038:
1034:
1027:
1023:
1019:
1015:
1010:
1003:
1001:
999:
995:
994:
989:
988:
983:
982:
975:
972:
963:
956:
954:
952:
951:
946:
945:
940:
939:
933:
928:
924:
923:
918:
909:
905:
902:
898:
893:
889:
884:
880:
875:
871:
867:
863:
862:glenoid fossa
859:
855:
846:
842:
839:
835:
831:
830:neural spines
827:
823:
819:
814:
810:
806:
797:
790:
788:
786:
782:
778:
774:
770:
766:
762:
758:
754:
746:
742:
737:
733:
730:
720:
716:
714:
710:
709:quadratojugal
705:
701:
697:
692:
688:
684:
680:
676:
675:
670:
669:
664:
663:
658:
655:The skull of
650:
645:
638:
636:
634:
630:
626:
625:frontal bones
622:
613:
605:
598:
593:
591:
589:
585:
581:
577:
573:
569:
564:
562:
558:
554:
550:
546:
542:
538:
534:
530:
526:
522:
518:
514:
511:, a complete
510:
506:
502:
498:
494:
490:
489:UMNH VP 20202
486:
481:
479:
475:
471:
467:
464:
460:
456:
452:
451:
447:
444:
440:
439:
435:
431:
427:
423:
419:
415:
407:
402:
395:
393:
391:
390:
385:
381:
377:
376:
371:
367:
363:
359:
355:
352:
348:
345:
341:
340:
327:
325:
316:
313:
312:Binomial name
309:
305:
304:
298:
295:
294:
290:
284:
283:
276:
273:
272:
269:
268:Ankylosaurini
263:
260:
259:
256:
250:
247:
246:
243:
237:
234:
233:
230:
224:
221:
218:
217:
214:
208:
205:
202:
201:
198:
192:
189:
186:
185:
182:
179:
176:
173:
172:
169:
166:
163:
162:
159:
156:
153:
152:
149:
146:
143:
142:
137:
132:
128:
123:
119:
114:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
39:
32:
26:
22:
19:
4210:
4125:
4118:
4111:
4104:
4097:
4090:
4083:
4076:
4071:Dyoplosaurus
4069:
4062:
4057:Ankylosaurus
4055:
4049:
4048:
4028:
4021:
4013:
4006:
3998:
3991:
3986:Pinacosaurus
3984:
3977:
3970:
3963:
3956:
3949:
3942:
3917:
3910:
3903:
3883:
3876:
3870:Hylaeosaurus
3868:
3861:
3854:
3846:
3838:
3806:
3799:
3794:Pawpawsaurus
3792:
3785:
3778:
3758:
3751:
3744:
3739:Denversaurus
3737:
3730:
3710:
3703:
3696:
3690:Peloroplites
3688:
3681:
3674:
3667:
3662:Borealopelta
3660:
3652:
3647:Acantholipan
3645:
3634:Nodosaurinae
3620:
3613:
3607:Peloroplites
3605:
3598:
3590:
3582:
3562:
3555:
3548:
3540:
3532:
3526:Mymoorapelta
3524:
3517:
3511:Hylaeosaurus
3509:
3502:
3495:
3487:
3479:
3472:
3467:Anoplosaurus
3465:
3458:
3447:Nodosauridae
3419:
3411:
3404:
3384:
3377:
3369:
3362:
3355:
3347:
3340:
3333:
3326:
3319:
3314:Cryptosaurus
3312:
3296:Ankylosauria
3274:Ankylosauria
3272:
3244:Ankylosauria
3232:Ornithischia
3217:Ornithischia
3169:
3165:Ornithischia
3160:
3151:
3123:Ankylosauria
3019:
3015:
3005:
2986:
2982:
2977:Leidyosuchus
2976:
2968:
2957:. Retrieved
2953:the original
2946:
2939:
2928:. Retrieved
2924:the original
2919:
2909:
2886:
2844:
2840:
2804:
2785:
2776:
2735:
2731:
2724:
2687:
2683:
2673:
2628:
2624:
2618:
2610:
2585:
2581:
2575:
2565:
2558:
2548:
2541:
2503:
2499:
2489:
2444:
2440:
2430:
2411:
2381:
2377:
2373:
2367:
2342:
2338:
2334:
2328:
2316:. Retrieved
2311:
2301:
2282:
2278:
2268:
2243:
2239:
2232:
2195:
2191:
2185:
2125:
2073:Leidyosuchus
2071:
2065:
2058:Helopanoplia
2056:
2047:
2038:
2028:
2018:
2012:
2006:
2002:Arvinachelys
2000:
1994:
1987:include the
1984:
1981:ornithomimid
1974:
1965:
1953:
1944:
1942:caenagnathid
1920:
1918:ankylosaurid
1908:
1898:
1894:Utahceratops
1892:
1876:
1875:hadrosaurid
1866:
1861:saurolophine
1856:
1855:
1819:Two Medicine
1815:Judith River
1802:
1801:
1795:
1673:Dyoplosaurus
1671:
1670:
1656:Ankylosaurus
1654:
1653:
1637:
1636:
1620:
1619:
1603:
1602:
1586:
1585:
1569:
1568:
1527:
1526:
1510:
1509:
1493:
1492:
1461:
1460:
1459:
1443:
1442:
1412:
1411:
1388:
1387:
1364:
1363:
1340:
1339:
1316:
1315:
1299:
1298:
1275:
1274:
1251:
1250:
1227:
1226:
1203:
1202:
1179:
1178:
1158:
1153:Ankylosaurus
1151:
1145:
1139:
1133:
1129:Pinacosaurus
1127:
1123:
1119:
1115:
1109:
1105:
1101:
1083:
1077:
1071:
1065:
1061:
1049:
1040:
1032:
1030:
1025:
1021:
1017:
1013:
997:
991:
985:
979:
976:
968:
948:
944:Ankylosaurus
942:
936:
922:Dyoplosaurus
920:
914:
896:
851:
802:
764:
750:
744:
740:
728:
725:
704:postorbitals
699:
695:
672:
668:Pinacosaurus
666:
660:
656:
654:
648:
618:
587:
583:
579:
571:
565:
488:
482:
455:alligatoroid
450:Arvinachelys
448:
436:
418:ankylosaurid
411:
387:
374:
373:
351:ankylosaurid
344:monospecific
338:
337:
336:
323:
318:
302:
301:
281:
280:
229:Ankylosauria
219:
203:
197:Ornithischia
187:
174:
24:
18:
4281:Ankylosaurs
4106:Scolosaurus
4023:Tsagantegia
3972:Jinyunpelta
3912:Shamosaurus
3863:Chuanqilong
3705:Silvisaurus
3683:Patagopelta
3615:Polacanthus
3564:Sauroplites
3364:Spicomellus
3342:Sarcolestes
3174:Thyreophora
2947:Deinosuchus
2335:Gryposaurus
2067:Deinosuchus
2014:Neurankylus
2008:Denazinemys
1907:ceratopsid
1890:ceratopsids
1864:hadrosaurid
1605:Scolosaurus
1253:Tsagantegia
1229:Shamosaurus
1135:Scolosaurus
1111:Tsagantegia
1096:to Western
998:Scolosaurus
993:Scolosaurus
743:(left) and
691:premaxillae
594:Description
438:Gryposaurus
434:hadrosaurid
426:Kane County
248:Subfamily:
213:Thyreophora
36:76.26
4270:Categories
4174:See also:
4099:Platypelta
4092:Oohkotokia
4030:Zaraapelta
3905:Gobisaurus
3856:Cedarpelta
3848:Aletopelta
3801:Stegopelta
3746:Edmontonia
3732:Animantarx
3698:Sauropelta
3676:Nodosaurus
3542:Priconodon
3386:Vectipelta
3321:Dracopelta
3253:see below↓
3156:Dinosauria
3022:: e10766.
2959:2016-03-27
2949:from Utah"
2930:2017-11-03
2246:: 104340.
2104:References
2078:azhdarchid
1951:troodontid
1935:orodromine
1931:ornithopod
1927:nodosaurid
1588:Oohkotokia
1318:Zaraapelta
1181:Gobisaurus
1090:Cenomanian
1085:Zaraapelta
1037:Laramidian
957:Osteoderms
927:osteoderms
888:acetabulum
791:Postcrania
773:surangular
757:pterygoids
531:, partial
501:predentary
487:specimen,
181:Dinosauria
4226:Q55656519
4113:Talarurus
3993:Saichania
3780:Europelta
3760:Texasetes
3622:Taohelong
3519:Invictarx
3421:Stegouros
3139:Kingdom:
3074:Dinosaurs
2983:PaleoBios
2690:: e5910.
2602:131302174
2535:full text
2359:220413406
2260:212423361
2198:: e5016.
2081:pterosaur
2049:Basilemys
2030:Compsemys
2020:Thescelus
1963:avisaurid
1946:Hagryphus
1823:Fruitland
1807:Campanian
1390:Saichania
1124:Saichania
1120:Talarurus
1073:Talarurus
1067:Saichania
1058:Campanian
987:Saichania
917:tail club
883:olecranon
822:synsacrum
753:quadrates
687:lacrimals
647:Skull of
553:osteoderm
513:tail club
505:vertebrae
503:, dorsal
497:mandibles
463:sandstone
380:Laramidia
366:Campanian
296:Species:
154:Kingdom:
148:Eukaryota
4240:55770783
4220:Wikidata
4186:Category
4176:Timeline
4120:Ziapelta
3592:Gastonia
3489:Gastonia
3147:Chordata
3145:Phylum:
3141:Animalia
3048:33552741
2869:83687743
2768:13261338
2760:21253683
2716:30479894
2665:21949721
2625:PLOS ONE
2530:23864598
2481:20877459
2441:PLOS ONE
2285:: 1–17.
2224:30065856
2140:Archived
2087:See also
1851:molluscs
1839:alluvial
1827:Kirtland
1622:Ziapelta
870:coracoid
854:scapulae
781:mandible
588:johnsoni
521:coracoid
517:scapulae
485:holotype
459:theropod
414:skeleton
354:dinosaur
235:Family:
168:Chordata
164:Phylum:
158:Animalia
144:Domain:
4008:Tarchia
4000:Shanxia
3060:Portals
3039:7825364
2849:Bibcode
2740:Bibcode
2707:6238772
2656:3176273
2633:Bibcode
2521:3730592
2472:2929175
2449:Bibcode
2390:1304144
2318:7 March
2215:6063217
2037:turtle
2027:turtle
2023:, the
1996:Boremys
1992:turtles
1967:Mirarce
1529:Shanxia
1079:Tarchia
1022:Tarchia
981:Tarchia
932:chevron
892:ischium
881:is the
874:humerus
809:centrum
777:dentary
769:alveoli
761:rostrum
713:maxilla
639:Cranium
584:kephalè
578:words "
568:generic
549:phalanx
525:humerus
515:, both
495:, both
474:CT scan
321:†
300:†
274:Genus:
261:Tribe:
105:↓
4253:377176
3046:
3036:
2897:
2867:
2811:
2766:
2758:
2714:
2704:
2663:
2653:
2600:
2528:
2518:
2479:
2469:
2418:
2388:
2357:
2258:
2222:
2212:
2132:
2061:, the
2052:, the
2043:, the
2040:Adocus
2035:adocid
2033:, the
1989:baenid
1970:, the
1958:, the
1949:, the
1933:, the
1913:, the
1903:, the
1885:, the
1871:, the
1829:, and
1102:et al.
1062:et al.
1024:, and
901:fibula
866:muscle
683:jugals
679:orbits
580:akaina
570:name,
545:fibula
446:turtle
443:baenid
416:of an
364:(late
3965:Datai
3328:Minmi
3170:Clade
3161:Clade
3152:Clade
3016:PeerJ
2989:(3).
2865:S2CID
2764:S2CID
2684:PeerJ
2598:S2CID
2386:JSTOR
2355:S2CID
2256:S2CID
2192:PeerJ
1955:Talos
834:ilium
576:Greek
541:tibia
537:femur
533:ilium
493:skull
347:genus
220:Clade
204:Clade
188:Clade
175:Clade
4127:Zuul
3230:see
3203:see
3086:Utah
3044:PMID
2895:ISBN
2809:ISBN
2756:PMID
2712:PMID
2661:PMID
2526:PMID
2477:PMID
2416:ISBN
2320:2022
2220:PMID
2130:ISBN
2070:and
2017:and
1897:and
1150:and
1118:and
1094:Asia
1082:and
1045:Asia
990:and
941:and
915:The
879:ulna
852:The
818:ribs
751:The
671:and
566:The
529:ulna
509:ribs
483:The
478:2018
430:Utah
384:2018
358:Utah
289:2018
44:Preęž’
4235:EoL
3034:PMC
3024:doi
2991:doi
2857:doi
2748:doi
2702:PMC
2692:doi
2651:PMC
2641:doi
2590:doi
2516:PMC
2508:doi
2504:280
2467:PMC
2457:doi
2347:doi
2287:doi
2248:doi
2244:108
2210:PMC
2200:doi
1841:to
559:in
349:of
4272::
4250::
4237::
4222::
3172::
3163::
3154::
3042:.
3032:.
3018:.
3014:.
2987:30
2985:.
2981:.
2918:.
2877:^
2863:.
2855:.
2845:35
2843:.
2839:.
2823:^
2793:^
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2762:.
2754:.
2746:.
2736:98
2734:.
2710:.
2700:.
2686:.
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2659:.
2649:.
2639:.
2627:.
2623:.
2596:.
2586:25
2584:.
2524:.
2514:.
2502:.
2498:.
2475:.
2465:.
2455:.
2443:.
2439:.
2398:^
2382:53
2380:.
2353:.
2343:26
2341:.
2283:10
2281:.
2277:.
2254:.
2242:.
2218:.
2208:.
2194:.
2190:.
2148:^
2138:.
2112:^
2083:.
2011:,
2005:,
1999:,
1825:,
1821:,
1817:,
1813:,
1156:.
1144:,
1138:,
1132:,
1126:,
1114:,
1076:,
1070:,
1020:,
984:,
635:.
563:.
547:,
543:,
539:,
535:,
527:,
523:,
519:,
499:,
428:,
424:,
392:.
222::
206::
190::
177::
94:Pg
38:Ma
33:,
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3625:?
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3374:?
3352:?
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3050:.
3026::
3020:9
2999:.
2993::
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2962:.
2933:.
2903:.
2871:.
2859::
2851::
2817:.
2770:.
2750::
2742::
2718:.
2694::
2688:6
2667:.
2643::
2635::
2629:6
2604:.
2592::
2532:.
2510::
2483:.
2459::
2451::
2445:5
2424:.
2392:.
2361:.
2349::
2322:.
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2289::
2262:.
2250::
2226:.
2202::
2196:6
747:.
278:â€
265:â€
252:â€
239:â€
226:â€
210:â€
194:â€
99:N
89:K
84:J
79:T
74:P
69:C
64:D
59:S
54:O
49:ęž’
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