Djadochta Formation

The Djadochta formation (sometimes transcribed and also known as Djadokhta, Djadokata, or Dzhadokhtskaya) is a highly fossiliferous geological formation in Central Asia, Gobi Desert, dating from the Late Cretaceous period, about 75 million to 71 million years ago. The type locality is the Bayn Dzak locality, famously known as the Flaming Cliffs. Reptile (including dinosaur) and mammal remains are among the fossils recovered from the formation.

Djadochta Formation
Stratigraphic range: Campanian, ~75–71 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Bayn Dzak (Flaming Cliffs), the type locality of the Djadochta Formation
Type	Geological formation
Unit of	Shamo Group
Sub-units	Bayn Dzak Member, Tugrugyin Member
Underlies	Barun Goyot Formation
Overlies	Alagteeg Formation
Area	Nemegt Basin, Ulan Nur Basin
Thickness	over 90 m (300 ft)
Lithology
Primary	Sandstone
Other	Mudstone
Type section
Named for	Shabarakh Usu (Bayn Dzak)
Named by	Berkey & Morris
Location	Flaming Cliffs
Year defined	1927
Coordinates	44°08′19″N 103°43′40″E
Approximate paleocoordinates	30.7°N 9.2°E / 30.7; 9.2
Region	Ömnögov
Country	Mongolia
Thickness at type section	about 90 m (300 ft)

Excavation history

Cretaceous-aged dinosaur fossil localities of Mongolia. Djadochta localities at area B.

The Djadochta formation was first documented and explored—though only a single locality—during paleontological expeditions of the American Museum of Natural History in 1922–1925, which were part of the Central Asiatic expeditions. The expeditions were led by Roy Chapman Andrews, in company of Walter Willis Granger as chief paleontologist and field team. The team did extensive exploration at the Bayn Dzak (formerly Shabarakh Usu) region, which they nicknamed flaming cliffs, given that at sunset the sediments of this locality had a characteristic reddish color. Notable finds included the first known fossils of Oviraptor, Protoceratops, Saurornithoides, and Velociraptor, the first confirmed dinosaur eggs (a partial nest of Oviraptor), as well as fossil mammals. Some of these were briefly described by Henry Fairfield Osborn during the ongoing years of the expeditions. In 1927 the formation was formally described and established by Berkey and Morris, with Bayn Dzak as the type locality.^[1][2]

In 1963 the Mongolian paleontologist Demberelyin Dashzeveg reported the discovery of a new fossiliferous locality of the Djadochta Formation: Tugriken Shireh.^[3] During the 1960s to 1970s, Polish-Mongolian and Russian-Mongolian paleontological expeditions collected new, partial to complete specimens of Protoceratops and Velociraptor at this locality, making these dinosaur species a common occurrence in Tugriken Shireh.^[4] Some of the most notable excavations made at Tugriken Shireh include the Fighting Dinosaurs (Protoceratops and Velociraptor locked in combat),^[5][6] and abundant articulated, in situ (in the original pose), and sometimes complete skeletons of Protoceratops.^[7][8]

During the 1980s a joint Soviet-Mongolian paleontological expedition discovered several Mesozoic fossil-rich localities in the Gobi Desert of Mongolia. Among these sites, Udyn Sayr was discovered and examined by the expedition, regarding its age as Late Cretaceous. This new locality was predominantly rich in avimimid fossils, with a lesser abundance of mammal and other dinosaur fossils.^[4]

In 1993 a collaborative expedition by the Mongolian Academy of Sciences and the American Museum of Natural History discovered a new fossil site within the Djadochta Formation, named Ukhaa Tolgod, which translates to "Brown Hills." This site has yielded a significant number of well-preserved fossils, including those of mammals, dinosaurs, lizards, and eggs. Most specimens are found in near-complete articulation, indicating excellent preservation conditions. Compared to other Mesozoic fossil sites, Ukhaa Tolgod stands out for its high fossil diversity.^[9][10]

Description

The modern-day Djadochta Formation is set in an arid habitat of sand dunes with little freshwater apart from oases and arroyos, in the Gobi Desert. The dominant lithology of the Djadochta Formation is represented by non-marine, cemented reddish-orange and pale orange to light gray, medium to fine-grained sands and sandstones, which include minor deposits of calcareous concretions and orange-brown silty clay. Less abundant sedimentation comprises conglomerates, siltstones, fluvial (water-deposited) sandstones, and mudstones. The entire thickness of the formation in the Ulan Nur Basin is at least 80 m (260 ft). Several aeolian processes (wind works) indicate the presence of large straight-crested dune-like structures, and smaller barchan (crescent-shaped) and parabolic (poorly U-shaped) dunes across the formation.^[11][12][10] Reddish sandstones are observed in numerous localities.^[12][10]

Exposures at the Flaming Cliffs

• Bayn Dzak (also spelled Bain Dzak, Bayanzag, Bayn Zag, Bayan Zag, or Shabarakh Usu; locally known as Flaming Cliffs): It is dominated by reddish-orange sandstones and well-sorted, unbedded, and medium-grained sands. The thickness of the strata at the Flaming Cliffs at least more than 30 m (98 ft). Less abundant lithology of Bayn Dzak includes cemented and poorly-cemented siltstones, mudstones, and grayish conglomerates. The latter are better exposed at western escarpments of the Flaming Cliffs.^[13][12] Bayn Dzak is about 90 m (300 ft) in total thickness and can be divided into two sections: alternations of horizontally-bedded sandstone and mudstone in the lowermost part, and sandstone-dominated successions in the upper or main part.^[14]
• Tugriken Shireh (also spelled Tugrik, Toogreeg, Toogreek, Tugreek, Tugrug, Turgrugyin, Tugrugeen, Tögrögiin, Tugrikiin, or Tugrikin): This locality is about 30 m (98 ft) in thickness and characterized by poorly cemented, fine-grained sandstones that have colors varying from pink to yellowish-white. The predominant mineral is quartz, and lesser common minerals are represented by feldspars and lithic fragments. Both cross-stratified and structureless sandstones are scattered across Tugriken Shireh.^[15][12]
• Udyn Sayr (also spelled Udan Sayr, Udan Sair, Ulaan Sair, or Üüden Sair): Sediments of this locality are exposed across a region of more than 60 km² (23 sq mi). It is divided into lower (thickness of at least more than 10 m (33 ft)) and upper (thickness of about 50 m (160 ft)) beds. The lower beds are fluvial originated and dominated by sandstones and mudstones. The upper beds are likely of aeolian origin and consist of reddish, cross-stratified and structureless sandstones.^[16]
• Ukhaa Tolgod (also spelled Oka Tolga): The strata exposed at Ukhaa Tolgod is dominated by reddish sandstones, with some sandstones containing small amounts of conglomeratic lenses and/or cobbles and pebbles. Conglomerate itself is in this site, and to a lesser level are mudstones and siltstones, which are thin and laterally restricted. Cross-stratified and fine-structured sandstones are particularly abundant at Ukhaa Tolgod.^[17][10]
• Zamyn Khondt (also spelled Dzamyn Khondt, Zamin Khond, or Dzamin Khond): This locality is characterized by reddish, well-sorted, and fine-grained sandstones with calcareous concretions. Some aeolian beds are present and are finely stratified to massive, having a visible thickness of about 20 m (66 ft).^[16]

Stratigraphy and age

The Djadochta formation occurs in the Late Cretaceous period of the Campanian stage. Magnetostratigraphic datings from the Bayn Dzak and Tugriken Shireh localities suggest that the Djadochta Formation was deposited during a time of rapidly changing polarity at about 75 million to 71 million years ago.^[12]

The Djadochta formation is separated into a lower Bayn Dzak Member and an upper Turgrugyin Member, which represent very similar depositional environments.^[12] Further strata from the Bayn Dzak Member includes that of the Ukhaa Tolgod locality, and its overall age is regarded also within the Campanian.^[10]

Based on the superposition of the members, the Tugrugyin Member overlies the Bayn Dzak Member making it somewhat younger, which indicates that the Bayn Dzak paleofauna lived somewhat earlier than that from Tugriken Shireh. However, it is not yet understood the precise temporal difference:^[12] Localities within the Djadochta Formation are considered to represent a sequence of progressively younger sediments and thereby paleofaunas. Ukhaa Tolgod may be younger than both Bayn Dzak and Tugriken Shireh.^[18] Based on their fossil record and strata, Udyn Sayr and Zamyn Khondt have been correlated with other Djadokhta localities, though fossils of Udyn Sayr may indicate that this locality is younger than Bayn Dzak and Tugriken Shireh.^[19]

Examinations on the strata of the Alag Teg (also spelled Alag Teeg or Alag Teer) locality, once considered part of this formation, indicates that it belongs to a different geological formation: the Alagteeg Formation, which is slightly older than the overlying Djadochta Formation. Based on sediments and stratigraphic relationships, the lower part of the Bayn Dzak locality is correlated with the Alag Teg locality, making both sections part of the Alagteeg Formation. The upper or main part of the former locality is considered part of the Djadochta Formation itself, as it shares similar lithology and stratigraphic relationships with Tugriken Shireh.^[14]

Stratigraphy of the Djadochta Formation^[12][14]

Formation	Time period	Member	Lithology	Thickness	Image
Barun Goyot	Early Maastrichtian		Poorly cemented, fine and medium-grained red to reddish-brown sandstones.	~110 m (360 ft)
Djadochta	Campanian
		Turgrugyin	Pale orange to light gray (sometimes yellowish-white) sands and sandstones.	30 m (98 ft)
		Bayn Dzak	Reddish-orange, crossbedded, and structureless sandstones, with minor deposits of brown siltstones and mudstones.	90 m (300 ft)
Alagteeg	Early Campanian Santonian	"lower Bayn Dzak"	Alternating reddish brown mudstone and horizontally laminated sandstone, with ripple cross laminations and rhizoliths.	~15 m (49 ft)

Depositional environment

Based on strata and rock facies (such as sandstones and caliche) of the formation and coeval units (Bayan Mandahu) it is currently agreed that sediments of the Djadochta Formation were deposited by wind activity in arid paleoenvironments comprising sand dunes with a warm semi-arid climate.^[20][12][14] Fluvial sedimentation at the Ukhaa Tolgod locality indicates the presence of short-lived water bodies during the times of the formation, which also contributed to its deposition.^[10]

Taphonomy

Examples of the Djadochta Formation preservation: articulated Citipati (top) and Protoceratops (bottom) specimens

A vast majority of articulated specimens from the Djadochta Formation are found in unstructured sandstones, indicating burial in situ by high-energy sand-bearing events. Some buried Protoceratops individuals are preserved in distinctive postures involving the body and head arched upwards, suggesting that the animals died in the process of trying to free themselves from the body of sand, where they eventually fossilized. As they were unable to escape burial, the sandy mass prevented carcasses from being scavenged by vertebrates. Most of these "buried" specimens are found with bite traces and large borings (tunnel-like holes made by small invertebrates) on bone joints areas and other surfaces, indicating that after death they were largely scavenged by invertebrates, such as skin beetles.^[21][22][23]

It has been suggested that the repeated occurrence of these feeding traces at limb joints may reflect that the responsible scavengers focused on collagen at the joint cartilage of dried dinosaur carcasses as a source of nitrogen, which was very low in the arid Djadochta Formation environments.^[24]

Examinations at the fossil preservation and sediments of Ukhaa Tolgod indicates that preserved animals were buried alive by catastrophic dune collapses. It is thought to have occurred when sand dunes became oversaturated with water resulting in their sudden downfall; heavy rainfall events likely acted as the triggering mechanism for this collapse.^[9][25][10] Examples from the Ukhaa Tolgod preservation include Citipati (brooding adults entombed atop nests and eggs);^[26][27] Khaan (a pair in close proximity likely killed by a single collapse event);^[28] and Saichangurvel (individual buried alive by a muddy dune).^[29]

Paleobiota of the Djadochta Formation

Articulated Protoceratops from Tugriken Shireh. This dinosaur is one of the most common occurrences in the Djadochta Formation

Among fossils, Protoceratops is extremely common in Djadochta localities. Bayn Dzak is reported as one of the localities with the highest concentration of Protoceratops fossils and has been noted as the "Protoceratops fauna".^[30] Adjacent to Bayn Dzak, at Tugriken Shireh, Protoceratops is also abundant.^[15] Other common dinosaur components of the paleofauna include Pinacosaurus and Velociraptor.^[11] Small vertebrates like lizards and mammals are rather abundant and diverse, with Adamisaurus and Kryptobaatar being the most abundant representatives.^[29][31][30] The paleofauna of the Djadochta Formation is very similar in composition to the nearby and coeval-regarded Bayan Mandahu Formation of Inner Mongolia. The two formations share many of the same genera, but differ in species. For instance, the most common mammal in Djadochta is Kryptobaatar dashzevegi, while in Bayan Mandahu it is the closely related K. mandahuensis. Similarly, the dinosaur fauna of Djadochta includes Protoceratops andrewsi and Velociraptor mongoliensis, which Bayan Mandahu yields P. hellenikorhinus and V. osmolskae.^[20][32]

Although fossil plants are extremely rare in the Djadochta Formation, the great abundancy of herbivorous Protoceratops at the arid-deposited Tugriken Shireh locality indicates that it had a moderate coverage of bushes or other low-growing plants.^[15]

The relatively low paleobiodiversity and climate settings of the Djadochta suggest that these conditions contributed to stressed paleoenvironments. Most of the fossil occurrences in the formation are occupied by Protoceratops, and small to medium-sized ankylosaurs, oviraptorids, and dromaeosaurids make much of the overall paleofauna. Large-bodied animals are absent or extremely rare in the formation. Comparisons with the Nemegt Formation further reflects stressed paleoenvironments. In contrast to Djadochta, Nemegt has yielded an extensive diversity of large dinosaur taxa, such as Deinocheirus, Nemegtosaurus, Saurolophus, Tarbosaurus, or Therizinosaurus. Most of these taxa are herbivorous, which combined with the mesic (well-watered) settings of the Nemegt Formation allowed the development of giant herbivores, in contrast to the stressed Djadochta Formation. Another indicative of stressed paleoenvironments is the almost non-existent amount of fully aquatic animals. Turtles are rarely recovered, and most are terrestrial such as Zangerlia.^[30] It is suggested that most of the fragmented hadrosaur, tyrannosaur and sauropod remains across the formation likely belong to non-endemic, passing by species.^[30]

Color key Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon		Notes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Flora

Genus	Species	Location	Material	Notes	Images
Radicites	R. gobiensis	Bayn Dzak	"Twenty plant roots."[33]	A tracheophyte, likely conifer.

Amphibians

Genus	Species	Location	Material	Notes	Images
Gobiates	Indeterminate	Udyn Sayr	"Partial skeleton with partial urostyle."[34]	A frog.

Crocodylomorphs

Genus	Species	Location	Material	Notes	Images
Artzosuchus	A. brachicephalus	Udyn Sayr	"Partial skull."[35]	A crocodylomorph.
Gobiosuchus	G. kielanae	Bayn Dzak	"Multiple specimens with partial skulls and skeletons."[36][37]	A gobisuchid.
	G.? parvus	Udyn Sayr	"Partial skull and skeleton."[38][39]	A gobisuchid.
Shamosuchus	S. djadochtaensis	Bayn Dzak, Ukhaa Tolgod	"Two skulls and partial skeleton."[40][41]	A paralligatorid.

Lizards

Genus	Species	Location	Material	Notes	Images
Adamisaurus	A. magnidentatus	Bayn Dzak, Tugriken Shireh, Ukhaa Tolgod	"Skulls and skeletons from multiple specimens."[42][29]	A teiid. Also present in the Barun Goyot Formation and Bayan Mandahu Formation.
Aiolosaurus	A. oriens	Ukhaa Tolgod	"Incomplete skull and partial skeleton."[29]	A varanoid.
Carusia	C. intermedia	Bayn Dzak, Ukhaa Tolgod	"Skulls from multiple specimens."[29]	A carusiid. Also present in the Barun Goyot Formation and Bayan Mandahu Formation.
Cherminotus	C. longifrons	Tugriken Shireh, Ukhaa Tolgod	"Skulls and partial skeleton."[29]	A varanoid. Also present in the Barun Goyot Formation.
Ctenomastax	C. parva	Zos	"Incomplete skull."[29]	An iguanid. Also present in the Barun Goyot Formation.
Dzhadochtosaurus	D. giganteus	Tugriken Shireh	"Partially complete skull."[43]	A macrocephalosaur.
Eoxanta	E. lacertifrons	Ukhaa Tolgod	"Incomplete skull."[29]	A scincomorph. Also present in the Barun Goyot Formation.
Estesia	E. mongoliensis	Bayn Dzak, Ukhaa Tolgod	"Partial skulls and teeth."[29]	A monstersaur. Also present in the Barun Goyot Formation.
Flaviagama	F. dzerzhinskii	Tugriken Shireh	"Skull and two vertebrae."[44]	A priscagamid.
Globaura	G. venusta	Bayn Dzak, Ukhaa Tolgod	"Partial skulls."[29]	A scincomorph. Also present in the Barun Goyot Formation.
Gobiderma	G. pulchrum	Udyn Sayr, Ukhaa Tolgod	"Skulls and skin impressions."[29]	A monstersaur. Also present in the Barun Goyot Formation.
Gobinatus	G. arenosus	Ukhaa Tolgod	"Partial skull."[29]	A teiid. Also present in the Barun Goyot Formation.
Hymenosaurus	H. clarki	Ukhaa Tolgod	"Partial skull."[29]	A scincomorph.
Isodontosaurus	I. gracilis	Bayn Dzak, Tugriken Shireh, Ukhaa Tolgod, Zos	"Numerous skulls and a partial skeleton."[29]	An iguanian. Also present in the Bayan Mandahu Formation.
Macrocephalosaurus	Indeterminate	Ukhaa Tolgod	"Partial skull and skeleton."[29]	A teiid. Also present in the Barun Goyot Formation.
Mimeosaurus	M. crassus	Bayn Dzak, Ukhaa Tolgod, Zos Wash	"Partially complete skulls."[29]	An acrodont. Also present in the Bayan Mandahu Formation.
Myrmecodaptria	M. microphagosa	Ukhaa Tolgod	"Single skull."[29]	A gekkotan.
Ovoo	O. gurval	Little Ukhaa Tolgod	"Partial skull."[45]	A varanid.
Parmeosaurus	P. scutatus	Ukhaa Tolgod	"Articulated skull and skeleton."[29]	A scincomorph.
Phrynosomimus	P. asper	Ukhaa Tolgod	"Two partial skulls."[29]	An acrodont. Also present in the Barun Goyot Formation.
Priscagama	P. gobiensis	Bayn Dzak, Ukhaa Tolgod	"Incomplete skulls."[29]	An priscagamid. Also present in the Barun Goyot Formation.
Saichangurvel	S. davidsoni	Ukhaa Tolgod	"Complete skull and skeleton in articulation."[46]	An iguanian.
Slavoia	S. darevskii	Ukhaa Tolgod	"Skulls and skeleton."[29]	A scincomorph. Also present in the Barun Goyot Formation.
Telmasaurus	T. grangeri	Bayn Dzak	"Partial skull and skeleton."[47][29]	A varanid.
Temujinia	T. ellisoni	Tugriken Shireh, Ukhaa Tolgod	"Several partial skulls."[29]	An iguanid.
Tchingisaurus	T. multivagus	Ukhaa Tolgod	"Partial skull."[29]	A teiid.
Unnamed scincomorph	Indeterminate	Ukhaa Tolgod	"Partial skull."[29]	A scincomorph.
Varanoidea indet.	Indeterminate	Ukhaa Tolgod	"Partial maxilla and vertebra."[29]	A varanoid.
Zapsosaurus	Z. sceliphros	Tugriken Shireh	"Two partial skulls."[29]	An iguanid.

Mammals

Genus	Species	Location	Material	Notes	Images
Asiatherium	A. reshetovi	Udyn Sayr	"Articulated skull and skeleton."[48]	A metatherian.
Bulganbaatar	B. nemegtbaataroides	Bayn Dzak, Ukhaa Tolgod	"Partial skull, and other remains."[49][9]	A multituberculate.
Catopsbaatar	C. catopsaloides	Ukhaa Tolgod	Not specified.[9]	A djadochtatheriid. Also present in the Barun Goyot Formation.
Chulsanbaatar	C. vulgaris	Ukhaa Tolgod	"Skull and partial skeleton."[9]	A multituberculate. Also present in the Barun Goyot Formation.
Deltatheridium	D. pretrituberculare	Bayn Dzak, Ukhaa Tolgod	"Partial skull and skeleton remains."[50][51]	A tribosphenid.
Deltatheroides	D. cretacicus	Bayn Dzak	"Partial skull."[50]	A djadochtatheriid.
Djadochtatherium	D. matthewi	Bayn Dzak, Tugriken Shireh	"Partial skulls."[52][53]	A djadochtatheriid.
Hyotheridium	H. dobsoni	Bayn Dzak	"Partial skull."[50]	A therian.
	Indeterminate	Ukhaa Tolgod	Not specified.[9]	A therian.
Kamptobaatar	K. kuczynskii	Bayn Dzak, Ukhaa Tolgod	"Partial skull, and other remains."[54][9]	A multituberculate.
Kennalestes	K. gobiensis	Bayn Dzak, Ukhaa Tolgod	"Nearly complete skull, and other remains."[55][9]	An eutherian.
Kryptobaatar	K. dashzevegi	Bayn Dzak, Tugriken Shireh, Ukhaa Tolgod	"Skulls and skeleton remains from several specimens."[54][31]	A djadochtatheriid. Gobibaatar and Tugrigbaatar are considered synonyms of this taxon.[56]
Maelestes	M. gobiensis	Ukhaa Tolgod	"Partial skull with skeleton."[57]	A cimolestid.
Mangasbaatar	M. udanii	Udyn Sayr	"Skulls and partial skeleton from two specimens."[58]	A djadochtatheriid.
Nemegtbaatar	N. gobiensis	Ukhaa Tolgod	Not specified.[9]	A multituberculate. Also present in the Barun Goyot Formation.
Sloanbaatar	S. mirabilis	Bayn Dzak, Ukhaa Tolgod	"Complete skull, and other remains."[54][9]	A multituberculate.
Tombaatar	T. sabuli	Ukhaa Tolgod	"Partial skull."[59]	A djadochtatheriid.
Ukhaatherium	U. nessovi	Ukhaa Tolgod	"Partial to nearly complete skeletons from several specimens."[60][61][62]	An eutherian.
Zalambdalestes	Z. lechei	Bayn Dzak, Tugriken Shireh	"Skulls and skeletons from several specimens."[50][63]	An eutherian.

Pterosaurs

Genus	Species	Location	Material	Notes	Images
Azhdarchidae indet.	Indeterminate	Tugriken Shireh	"Indeterminate bone inside the gut cavity of a Velociraptor."[64]	An azhdarchid.

Turtles

Genus	Species	Location	Material	Notes	Images
Nanhsiungchelyidae indet.	Indeterminate	Abdrant Nuru	"Three shell fragments."[65]	A nanhsiungchelyid.
	Indeterminate	Bayn Dzak	"Partial shells."[65]	A nanhsiungchelyid.
	Indeterminate	Udyn Sayr	"Two shell fragments."[65]	A nanhsiungchelyid.
Zangerlia	Z. dzamynchondi	Zamyn Khondt	"Partial shell."[66]	A nanhsiungchelyid.
	Z. ukhaachelys	Ukhaa Tolgod	"Partial skull and skeleton."[67]	A nanhsiungchelyid.

Dinosaurs

Ornithischians

Ankylosaurids

Genus	Species	Location	Material	Notes	Images
Minotaurasaurus	M. ramachandrani	Ukhaa Tolgod	[Two] complete skulls, mandibles, and first cervical half-ring.[68][69]	An ankylosaurid previously thought to be a junior synonym of Tarchia, but is now considered to be a valid and distinct taxon.
Pinacosaurus	P. grangeri	Bayn Dzak, Ukhaa Tolgod	[Three] skulls, mandibles, predentary, cervical vertebrae, dorsal vertebrae, caudal vertebrae, ribs, scapula, coracoids, humerus, radius, ulna, ilium, femora, tibia, fibula, pelvis, manus, tail club handles, cervical half-rings, osteoderms, and a nearly complete skeleton lacking a skull.[70][71][72]	An ankylosaurid also known from the Alagteeg Formation and Bayan Mandahu Formation.
Ankylosauridae indet.	Indeterminate	Zamyn Khondt	Partially complete postcranial skeleton with in situ osteoderms.[73]	Previously referred to Saichania, but is now referred to as Ankylosauridae indet., or cf. Pinacosaurus.[73]

Ceratopsians

Genus	Species	Location	Material	Notes	Images
Bainoceratops	B. efremovi	Bayn Dzak	"Partial vertebrae."[74]	A protoceratopsid. May be synonymous with Protoceratops.[75]
Bagaceratops	Indeterminate	Udyn Sayr	"Skull with partial skeleton."[19]	A protoceratopsid. Indeterminate between Bagaceratops and Protoceratops.[19]
Protoceratops	P. andrewsi	Bayn Dzak, Tugriken Shireh, Udyn Sayr, Zamyn Khondt	"Multiple partial to complete specimens."[76][7][77][8][19]	A protoceratopsid.
	P. hellenikorhinus	Bor Tolgoi, Udyn Sayr	"Partial cranial remains."[78]	A protoceratopsid.
Protoceratopsidae indet.	Indeterminate	Ukhaa Tolgod	"Numerous skulls and remains."[79][80]	A protoceratopsid.
Udanoceratops	U. tschizhovi	Udyn Sayr	"Skull and fragmented skeleton elements."[81]	A giant leptoceratopsid.

Hadrosaurs

Genus	Species	Location	Material	Notes	Images
Hadrosauroidea indet.	Indeterminate	Tugriken Shireh	"Fragmented remains from juveniles."[82][83]	A hadrosauroid.
Plesiohadros	P. djadokhtaensis	Alag Teeg	"Skull and partial body elements."[83]	A hadrosauroid. Actually hails from the Alagteeg Formation.[84]

Pachycephalosaurs

Genus	Species	Location	Material	Notes	Images
Goyocephale	G. lattimorei	Boro Khovil	"Partial skull and skeleton."[85]	A pachycephalosaurid. Locality sediments may belong to this formation.[85][86]

Saurischians

Sauropods

Genus	Species	Location	Material	Notes	Images
Sauropoda indet.	Indeterminate	Not specified.[87]	Not specified.[87]	A sauropod.

Theropods

Alvarezsaurs

Genus	Species	Location	Material	Notes	Images
Alvarezsauridae indet.	Indeterminate	Tugriken Shireh	"Partial skull, braincase, and skeleton elements of two specimens."[88][89]	An alvarezsaurid also known as the Tugriken Shireh alvarezsaur. Uncertainly referred as Parvicursor sp.[89]
Kol	K. ghuva	Ukhaa Tolgod	"A well-preserved right foot."[90]	A large alvarezsaurid. Its classification has been criticized.[91]
Shuvuuia	S. deserti	Ukhaa Tolgod	"Multiple specimens with skull and skeletons."[92][93][94][90][95]	An alvarezsaurid.
Undescribed Alvarezsauridae	Indeterminate	Bayn Dzak	"Partial pelvic girdle and hindlimb."[96]	An alvarezsaurid.
	Indeterminate	Gilvent Wash	Not given.[90]	An alvarezsaurid.

Birds

Genus	Species	Location	Material	Notes	Images
Apsaravis	A. ukhaana	Ukhaa Tolgod	"Partial postcranial skeleton."[97]	A basal ornithurine bird.
Elsornis	E. keni	Tugriken Shireh	"Partial articulated skeleton lacking the skull."[98]	An enantiornithe.
Gobipteryx	G. minuta	Ukhaa Tolgod	"Partial skull."[99]	An enantiornithine. Also present in the Barun Goyot Formation.
Protoceratopsidovum'	P. fluxuosum	Bayn Dzak	"Partial eggs."[100]	Eggs probably laid by a bird.[101]
	P. minimum	Baga Tariach, Tugriken Shireh	"Clutch of four eggs and one pole of egg."[100]	Eggs probably laid by a bird.[101]
	P. sincerum	Bayn Dzak, Tugriken Shireh	"Multiple eggs and shells."[100]	Eggs probably laid by a bird.[101]
Styloolithus	S. sabathi	Bayn Dzak	"Partial to complete eggs."[101]	Eggs probably laid by a bird.

Dromaeosaurs

Genus	Species	Location	Material	Notes	Images
Tsaagan	T. mangas	Ukhaa Tolgod	"Skull and partial skeleton."[102]	A dromaeosaurid
Velociraptor	V. mongoliensis	Bayn Dzak, Chimney Buttes, Gilvent Wash, Tugriken Shireh, Udyn Sayr, Ukhaa Tolgod	"Multiple partial to complete specimens."[103][104][105][106][107][108]	A dromaeosaurid.
Undescribed Dromaeosauridae	Indeterminate	Abdrant Nuru	"Claw."[109]	A dromaeosaurid.
	Indeterminate	Zos Wash	"Frontal region."[102]	A dromaeosaurid. Differs from Tsaagan.[102]

Halszkaraptorines

Genus	Species	Location	Material	Notes	Images
Halszkaraptor	H. escuilliei	Ukhaa Tolgod	"Partial skeleton with complete skull."[110]	A halszkaraptorine.
Mahakala	M. omnogovae	Tugriken Shireh	"Fragmented skull and skeleton."[111]	A halszkaraptorine.

Ornithomimosaurs

Genus	Species	Location	Material	Notes	Images
Aepyornithomimus	A. tugrikinensis	Tugriken Shireh	"Nearly complete foot."[112]	An ornithomimid.
Ornithomimosauria indet.	Indeterminate	Ukhaa Tolgod	"Partial braincase, jaw tips, ribs, and vertebral fragments".[113][114]	An ornithomimid.

Oviraptorosaurs

Genus	Species	Location	Material	Notes	Images
Avimimus	A. portentonsus	Udyn Sayr	"Partial skull and skeleton."[115][116]	An avimimid. Its locality origin has been disputed and may hail from the Nemegt Formation.[117]
Citipati	C. osmolskae	Ukhaa Tolgod	"Multiple specimens with partial to nearly complete skeletons, an embryo, eggs and nesting individuals."[118][119][120][27]	An oviraptorid.
Elongatoolithus	E. frustrabilis
	E. subtitectorius
Khaan	K. mckennai	Ukhaa Tolgod	"Several specimens with partial to complete skeletons and skulls."[119][121][28]	An oviraptorid.
Macroolithus	M. mutabilis		"Eggs."	Eggs probably laid by an oviraptorid
Oviraptor	O. philoceratops	Bayn Dzak	"Partial skeleton with skull, associated with a nest and juvenile."[122][120][27]	An oviraptorid.
Oviraptoridae indet.	Indeterminate	Zamyn Khondt	"Nearly complete skeleton with skull."[123]	An oviraptorid also known as the Zamyn Khondt oviraptorid. Uncertainly referred to Citipati.[119][120]
	Indeterminate	Zamyn Khondt	"Nearly complete skull with atlas and axis."[124]	An oviraptorid.
	Indeterminate	Udyn Sayr	"Assemblage of individuals."[27]	An oviraptorid.
	Indeterminate	Not specified.	"Two skulls with characteristic high crest."[125][126][127]	An oviraptorid.

Troodontids

Genus	Species	Location	Material	Notes	Images
Almas	A. ukhaa	Ukhaa Tolgod	"Skull with partial skeleton."[128]	A troodontid.
Archaeornithoides	A. deinosauriscus	Bayn Dzak	"Partial skull."[129]	A troodontid? Uncertain relationships among coelurosaurs.[129][130]
Byronosaurus	B. jaffei	Ukhaa Tolgod	"Skull and fragmentary skeleton."[131]	A troodontid.
Gobivenator	G. mongoliensis	Zamyn Khondt	"Almost complete skeleton."[132]	A troodontid.
Saurornithoides	S. mongoliensis	Bayn Dzak	"Skull with fragmentary skeleton."[133]	A troodontid.
Troodontidae indet.	Indeterminate	Ukhaa Tolgod	"Partial skeleton."[134]	A troodontid.
	Indeterminate	Ukhaa Tolgod	"Juvenile skulls, skeleton, and one nest."[135][130]	A troodontid. Referred to either Almas,[136][128] or Byronosaurus.[130]
	Indeterminate	Ukhaa Tolgod	"Fragmented skull and skeleton remains."[137]	A troodontid. Provisionally referred to Saurornithoides,[137] but now excluded.[133]
	Indeterminate	Ukhaa Tolgod	"Partial skull and skeletons from two specimens."[93]	A troodontid.

Tyrannosaurids

Genus	Species	Location	Material	Notes	Images
Tyrannosauridae indet.	Indeterminate	Bayn Dzak	Not specified.[32]	A tyrannosaurid.
	Indeterminate	Khongil	"Supraorbital, vertebra, rib, femur and metatarsals."[138]	A tyrannosaurid.
	Indeterminate	Not specified.	"Partial right ilium."[139]	A tyrannosaurid.
	Indeterminate	Not specified.	"Teeth."[20]	A tyrannosaurid.

Gallery

Panoramic view of the Flaming Cliffs (Bayn Dzak), type locality of the Djadochta Formation

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See also

• List of dinosaur-bearing rock formations
• List of fossil sites (with link directory)
• Barun Goyot Formation – Geological formation in Mongolia
• Bayan Mandahu Formation – Geological formation in China
• Nemegt Formation – Geological formation in Mongolia

References

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