Citipati

Citipati (pronounced [ˈtʃiːt̪ɪpət̪i] in Hindi, meaning 'funeral pyre lord') is a genus of oviraptorid theropod dinosaur from the Late Cretaceous Period of what is now Mongolia (specifically, the Djadokhta Formation of Ukhaa Tolgod, in the Gobi Desert). It is one of the best-known oviraptorids, thanks to a number of well-preserved skeletons, including several specimens found in brooding positions atop nests of eggs. These nesting specimens have helped to solidify the link between non-avian dinosaurs and birds.

The type species, Citipati osmolskae, was described in 2001. A second, as yet unnamed species may also exist. Citipati is often confused with the similar Oviraptor.

Citipati
Temporal range: Late Cretaceous, 84–75 Ma
Citipati IGM 100 979
Nesting C. osmolskae specimen nicknamed "Big Mamma", housed at the AMNH
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Oviraptoridae
Genus: Citipati
Clark, Norell & Barsbold, 2001
Species:
C. osmolskae
Binomial name
Citipati osmolskae
Clark, Norell & Barsbold, 2001

Description

Citipati profiles1
Artist's impression of C. osmolskae and the unnamed Citipati species

The largest Citipati were emu-sized animals and, at about 3 meters (10 ft) long, were the largest known oviraptorids until Gigantoraptor was described in 2007. Like other oviraptorids, Citipati had an unusually long neck and shortened tail, compared to most other theropods. Its skull was unusually short and highly pneumatized (riddled with openings in the bone structure), ending in a stout, toothless beak. Perhaps the most distinctive feature of Citipati was its tall crest, superficially similar to that of a modern cassowary. The crest was relatively low in the type species, C. osmolskae, with a nearly vertical front margin grading into the beak. In contrast, the crest of one referred specimen which has not yet been assigned a specific name (provisionally labeled C. sp.) was taller, with a prominent notch in the front margin, creating a squared appearance.

History of discovery

The type species of Citipati, C. osmolskae, was named by James M. Clark, Mark Norell, and Rinchen Barsbold in 2001, in honor of Halszka Osmólska, a noted paleontologist whose work has dealt extensively with oviraptorids and other Mongolian theropods. The name Citipati is formed from the Sanskrit words citi, meaning 'funeral pyre' and pati meaning 'lord' (चितिपति). In Tibetan Buddhist folklore, the Citipati were two monks who were beheaded by a thief, while deep in a meditative trance. The Citipati are often depicted as a pair of dancing skeletons surrounded by flame, hence the application of the name to the beautifully preserved oviraptorid skeletons.[1]

Citpatibcn1
Specimen GIN 100/42 which most popular restorations of Oviraptor are based on

Since the type skull of Oviraptor is so poorly preserved and crushed, another oviraptorid specimen (IGM or GIN 100/42) had become the quintessential depiction of that dinosaur, even appearing in scientific papers with the label Oviraptor philoceratops.[2] However, this distinctive-looking, tall-crested species has more features of the skull in common with Citipati than it does with Oviraptor and it may represent a second species of Citipati or possibly an entirely new genus, pending further study.[1] Additionally, the nesting oviraptorid specimens had gained widespread attention before they were actually referred to Citipati. While usually labeled simply "oviraptorids", they have on occasion been confused with Oviraptor itself. The fact that the first Oviraptor specimen was found on a nest as well confused the matter further. As it stands, most popular illustrations of Oviraptor actually depict Citipati and the present material available for Oviraptor itself is too fragmentary to be reliably reconstructed.

Classification

Citipati osmolskae
Skeletal restorations of C. osmolskae specimens
Zamyn Khondt Citipati
Skeletal restorations of GIN 100/42

The cladogram below follows an analysis by Fanti et al., 2012.[3]

Oviraptoridae

Oviraptor

unnamed

Rinchenia

Citipati

unnamed

Khaan

unnamed

Conchoraptor

unnamed

Machairasaurus

unnamed

"Ingenia" (=Ajancingenia)

unnamed

Nemegtomaia

Heyuannia

Paleobiology

Reproduction

Nesting Citipati
Nesting specimen nicknamed "Big Auntie"

At least four Citipati specimens have been found in brooding positions, the most famous of which, a large specimen nicknamed "Big Mamma", was first announced (but not named) in 1995,[4] described in 1999,[5] and referred to the genus Citipati in 2001.[1] All of the nesting specimens are situated on top of egg clutches, with their limbs spread symmetrically on each side of the nest, front limbs covering the nest perimeter. This brooding posture is found today only in birds and supports a behavioral link between birds and theropod dinosaurs.[5] The nesting position of Citipati also supports the hypothesis that it and other oviraptorids had feathered forelimbs. With the 'arms' spread along the periphery of the nest, a majority of eggs would not be covered by the animal's body unless an extensive coat of feathers was present.[6]

Although fossilized dinosaur eggs are rare, Citipati eggs and oviraptorid eggs in general, are relatively well known. Along with the four known nesting specimens, dozens of isolated oviraptorid nests have been uncovered in the Gobi Desert. Citipati eggs are shaped like elongated ovals (elongatoolithid) and resemble the eggs of ratites in texture and shell structure. In the nest, Citipati eggs are typically arranged in concentric circles of up to three layers, and a complete clutch may have consisted of as many of 22 eggs.[7] The eggs of Citipati are the largest known definitive oviraptorid eggs, at 18 cm. In contrast, eggs associated with Oviraptor are only up to 14 cm long.[5]

Citipati egg
C. osmolskae egg with preserved embryo

Ironically, it was the association with eggs that gave oviraptorids their name (which means 'egg thieves'). The first oviraptorid eggs (of the genus Oviraptor) were found in close proximity to the remains of the ceratopsian dinosaur Protoceratops and it was assumed that the oviraptorids were preying upon the eggs of the ceratopsians.[8] It was not until 1993, when a Citipati embryo was discovered inside an egg of the type assigned to Protoceratops, that the error was corrected.[9] Norell et al., who recognized the embryo as oviraptorid, assigned it to the genus Citipati in 2001, based on the vertical orientation of the premaxilla (a bone structure at the tip of the jaw), a feature found only in Citipati. The egg containing the embryo was smaller than most known Citipati eggs at only 12 cm, though it was partially eroded and broken into three pieces, making an accurate estimate of its original size difficult.[5] The embryo-bearing egg was otherwise identical to other oviraptorid eggs in shell structure and was found in an isolated nest, again arranged in a circular pattern.[9]

Two skulls belonging to very young or embryonic Byronosaurus were found associated with the same nest as the first Citipati embryo.[10] It is possible that these tiny troodontids were preyed upon by the Citipati. Alternately, Mark Norell suggested that the juvenile troodonts were raiding the Citipati nest, or even that an adult Byronosaurus had laid eggs in a Citipati nest as an act of nest parasitism.[9]

See also

References

  1. ^ a b c Clark, J.M., Norell, M.A., & Barsbold, R. (2001). "Two new oviraptorids (Theropoda:Oviraptorosauria), upper Cretaceous Djadokhta Formation, Ukhaa Tolgod, Mongolia." Journal of Vertebrate Paleontology 21(2):209-213., June 2001.
  2. ^ Barsbold, R., Maryanska, T., and Osmolska, H. (1990). "Oviraptorosauria," in Weishampel, D.B., Dodson, P., and Osmolska, H. (eds.). The Dinosauria. Berkeley: University of California Press, pp. 249-258.
  3. ^ Fanti F, Currie PJ, Badamgarav D (2012). "New Specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia." PLoS ONE, 7(2): e31330. doi:10.1371/journal.pone.0031330
  4. ^ Norell, M.A., Clark, J.M., Chiappe, L.M., and Dashzeveg, D. (1995). "A nesting dinosaur." Nature 378:774-776.
  5. ^ a b c d Clark, J.M., Norell, M.A., & Chiappe, L.M. (1999). "An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avianlike brooding position over an oviraptorid nest." American Museum Novitates, 3265: 36 pp., 15 figs.; (American Museum of Natural History) New York. (5.4.1999).
  6. ^ Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
  7. ^ Varricchio, D.J. (2000). "Reproduction and Parenting," in Paul, G.S. (ed.). The Scientific American Book of Dinosaurs. New York: St. Martin's Press, pp. 279-293.
  8. ^ Osborn, H.F. (1924). "Three new Theropoda, Protoceratops zone, central Mongolia." American Museum Novitates, 144: 12 pp., 8 figs.; (American Museum of Natural History) New York. (11.7.1924).
  9. ^ a b c Norell, M. A., J. M. Clark, D. Dashzeveg, T. Barsbold, L. M. Chiappe, A. R. Davidson, M. C. McKenna, and M. J. Novacek (1994). "A theropod dinosaur embryo, and the affinities of the Flaming Cliffs Dinosaur eggs." Science 266: 779–782.
  10. ^ Bever, G.S. and Norell, M.A. (2009). "The perinate skull of Byronosaurus (Troodontidae) with observations on the cranial ontogeny of paravian theropods." American Museum Novitates, 3657: 51 pp.

External links

Caenagnathoidea

Caenagnathoidea ("recent jaw forms") is a group of advanced oviraptorosaurian dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, often with bony crests atop the head. They ranged in size from Caudipteryx, which was the size of a turkey, to the 8 meter long, 1.4 ton Gigantoraptor. The group (along with all maniraptoran dinosaurs) is close to the ancestry of birds. The most complete specimens have been found in Asia, representing members of the sub-group Oviraptorinae. Notable but fragmentary remains are also known from North America, almost all of which belong to the subgroup Elmisaurinae.The earliest definitive caenangnathoid is Microvenator celer, which dates to the late Aptian age of the early Cretaceous period, though the slightly earlier Caudipteryx from the lower Yixian Formation of China, may also be a member of this group.

Campanian

The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous series. Campanian spans the time from 83.6 (± 0.7) to 72.1 (± 0.6) million years ago. It is preceded by the Santonian and it is followed by the Maastrichtian.The Campanian was an age when a worldwide sea level rise covered many coastal areas. The morphology of some of these areas has been preserved: it is an unconformity beneath a cover of marine sedimentary rocks.

Citipati (Buddhism)

Citipati(Sanskrit: चितिपति) is a protector deity or supernatural being in Tibetan Buddhism and Vajrayana Buddhism of India. It is formed of two skeletal deities, one male and the other female, both dancing wildly with their limbs intertwined inside a halo of flames representing change. The Citipati is said to be one of the seventy-five forms of Mahakala. Their symbol is meant to represent both the eternal dance of death as well as perfect awareness. They are invoked as ‘wrathful deities’, benevolent protectors or fierce beings of demonic appearance. The dance of the Citipati is commemorated twice annually in Tibet.

Corythoraptor

Corythoraptor (meaning "crested raptor") is a genus of crested oviraptorid theropod dinosaur from the Nanxiong Formation of China. It is known from one species, C. jacobsi, named after palaeontologist Louis L. Jacobs. Including it, there are seven oviraptorids known from the Nanxiong Formation, showing a high level of diversity in the area, and that the different taxa may have occupied different ecological niches.

Dharmapala

A dharmapāla (Wylie: chos skyong) is a type of wrathful god in Buddhism. The name means "Dharma protector or defender" in Sanskrit, and the dharmapālas are also known as the Defenders of the Law (Dharma), or the Protectors of the Law.

Elongatoolithidae

Elongatoolithidae is an oofamily of fossil eggs, representing the eggs of oviraptorosaurs (with the exception of the avian Ornitholithus). They are known for their highly elongated shape. Elongatoolithids have been found in Europe, Asia, and both North and South America.

Guhyasamāja Tantra

The Guhyasamāja Tantra (Sanskrit: Guhyasamājatantra; Tibetan: Gsang ’dus rtsa rgyud (Toh 442); Tantra of the Secret Community) is one of the most important scriptures of Tantric Buddhism. In its fullest form, it consists of seventeen chapters, though a separate "explanatory tantra" (vyākhyātantra) known as the Later Tantra (Sanskrit: Guhyasamāja Uttaratantra; Tibetan: Rgyud phyi ma. (Toh 443)) is sometimes considered to be its eighteenth chapter. Many scholars believe that the original core of the work consisted of the first twelve chapters, with chapters thirteen to seventeen being added later as explanatory material.

In India, it was classified as a Yoga or Mahāyoga Tantra. In Tibet it is considered an Unexcelled Yoga Tantra (rnal ’byor bla med rgyud). It develops traditions found in earlier scriptures such as the Compendium of Reality (Sanskrit: Sarva-tathāgata-tattva-saṃgraha; De bzhin gshegs pa thams

cad kyi de kho na nyid bsdus pa (Toh 479)) but is focused to a greater extent on the antinomian aspects characteristic of the later Buddhist Tantras. Naropa and Aryadeva considered the Compendium of Reality to be a root tantra in relation to the Guhyasamaja Tantra. The Guhyasamaja Tantra survives in Sanskrit manuscripts and in Tibetan and Chinese translation.

The Guhyasiddhi of Padmavajra, a work associated with the Guhyasamaja tradition, prescribes acting as a Saiva guru and initiating members into Shaiva Siddhanta scriptures and mandalas. Statues and pictures were also created to beautify the radical methodology. However, because extraordinary sexual acts with women including many virgins had no integrity to fundamental buddhist precepts, it is no longer done in Tibetan Buddhism. On the contrary, warning symbols and festivals, such as Citipati, are widely spread.

Heyuannia

Heyuannia ("from Heyuan") is a genus of oviraptorid dinosaur that lived during the Late Cretaceous Period in China. It was the first oviraptorid found in that country; most others were found in neighbouring Mongolia. Two species are known: H. huangi, named by Lü Junchang in 2002; and H. yanshini, originally named as a separate genus Ingenia by Rinchen Barsbold in 1981 (and renamed to Ajancingenia in 2013 due to the preoccupation of Ingenia).

Huanansaurus

Huanansaurus is an extinct genus of oviraptorid dinosaur that lived approximately 72 million years ago, between the Campanian and Maastrichtian, during the latter part of the Cretaceous period in what is now China, in the Nanxiong Formation.

Khaan

Khaan (; Mongol [χaːŋ] 'lord') was an oviraptorid dinosaur that was found in the Djadochta Formation of Mongolia and lived in the Late Cretaceous Period (Campanian), 75 million years ago.

Mahakala

Mahakala (Sanskrit: महाकाल; IAST: Mahākāla) is a deity common to Hinduism, Buddhism and Sikhism. According to Hinduism, Mahakala is a manifestation of Shiva and is the consort of Hindu Goddess Kali and most prominently appears in Kalikula sect of Shaktism. Mahākāla also appears as a protector deity known as a dharmapala in Vajrayana Buddhism, particularly most Tibetan traditions (Citipati), in Tangmi (Chinese Esoteric Buddhism) and in Shingon (Japanese Esoteric Buddhism). He is known as Dàhēitiān and Daaih'hāktīn (大黑天) in Mandarin and Cantonese, Daeheukcheon (대흑천) in Korean and Daikokuten (大黒天) in Japanese. In Sikhism, Mahākāla is referred to as Kal, who is the governor of Maya.

Mark Norell

Mark A. Norell (born July 26, 1957) is an American paleontologist and molecular geneticist, acknowledged as one of the most important living vertebrate paleontologists. He is currently the chairman of paleontology and a research associate at the American Museum of Natural History. He is best known as the discoverer of the first theropod embryo and for the description of feathered dinosaurs. Norell is credited with the naming of the genera Apsaravis, Byronosaurus, Citipati, Tsaagan, and Achillobator. His work regularly appears in major scientific journals (including cover stories in Science and Nature) and was listed by Time magazine as one of the ten most significant science stories of 1993, 1994 and 1996.

Norell is both a fellow of the Explorer's Club and the Willi Hennig Society.

Memento mori

Memento mori (Latin: "remember (that) you will die") is the medieval Latin Christian theory and practice of reflection on mortality, especially as a means of considering the vanity of earthly life and the transient nature of all earthly goods and pursuits. It is related to the ars moriendi ("The Art of Dying") and similar Western literature. Memento mori has been an important part of ascetic disciplines as a means of perfecting the character by cultivating detachment and other virtues, and by turning the attention towards the immortality of the soul and the afterlife.In art, memento mori are artistic or symbolic reminders of mortality. In the European Christian art context, "the expression [...] developed with the growth of Christianity, which emphasized Heaven, Hell, and salvation of the soul in the afterlife".

Nankangia

Nankangia is an extinct genus of caenagnathoid oviraptorosaurian dinosaur known from the Late Cretaceous Nanxiong Formation of Nankang County, Ganzhou City of Jiangxi Province, southeastern China. It contains a single species, Nankangia jiangxiensis. N. jiangxiensis coexisted with at least four other caenagnathoids, including an unnamed oviraptorid, Banji long, Ganzhousaurus nankangensis and Jiangxisaurus ganzhouensis. The relatively short dentary and non-downturned mandibular symphysis of Nankangia suggest that it may have been more herbivorous than carnivorous.

Nemegtomaia

Nemegtomaia is a genus of oviraptorid dinosaur from what is now Mongolia that lived in the Late Cretaceous Period, about 70 million years ago. The first specimen was found in 1996, and became the basis of the new genus and species N. barsboldi in 2004. The original genus name was Nemegtia, but this was changed to Nemegtomaia in 2005, as the former name was preoccupied. The first part of the generic name refers to the Nemegt Basin, where the animal was found, and the second part means "good mother", in reference to the fact that oviraptorids are known to have brooded their eggs. The specific name honours the palaeontologist Rinchen Barsbold. Two more specimens were found in 2007, one of which was found on top of a nest with eggs, but the dinosaur had received its genus name before it was found associated with eggs.

Nemegtomaia is estimated to have been around 2 m (7 ft) in length, and to have weighed 40 kg (85 lb). As an oviraptorosaur, it would have been feathered. It had a deep, narrow, and short skull, with an arched crest. It was toothless, had a short snout with a parrot-like beak, and a pair of tooth-like projections on its palate. It had three fingers; the first was largest and bore a strong claw. Nemegtomaia is classified as a member of the oviraptorid subfamily Ingeniinae, and is the only known member of this group with a cranial crest. Though Nemegtomaia has been used to suggest that oviraptorosaurs were flightless birds, the clade is generally considered a group of non-avian dinosaurs.

The nesting Nemegtomaia specimen was placed on top of what was probably a ring of eggs, with its arms folded across them. None of the eggs are complete, but they are estimated to have been 5 to 6 cm (2 to 2.3 in) wide and 14 to 16 cm (5 to 6 in) long when intact. The specimen was found in a stratigraphic area that indicates Nemegtomaia preferred nesting near streams that would provide soft, sandy substrate and food. Nemegtomaia may have protected its eggs by covering them with its tail and wing feathers. The skeleton of the nesting specimen has damage that indicates it was scavenged by skin beetles. The diet of oviraptorids is uncertain, but their skulls are most similar to other animals that are known or thought to have been herbivorous. Nemegtomaia is known from the Nemegt and Baruungoyot formations, which are thought to represent humid and arid environments that coexisted in the same area,

Oviraptor

Oviraptor is a genus of small Mongolian theropod dinosaurs, first discovered by technician George Olsen in an expedition led by Roy Chapman Andrews, and first described by Henry Fairfield Osborn, in 1924. Its name is Latin for 'egg taker' or "egg seizer", referring to the fact that the first fossil specimen was discovered atop a pile of what were thought to be Protoceratops eggs, and the specific name philoceratops means "lover of ceratopsians", also given as a result of this find. In his 1924 paper, Osborn explained that the name was given due to the close proximity of the skull of Oviraptor to the nest (it was separated from the eggs by only 4 inches or 10 centimetres of sand). However, Osborn also suggested that the name Oviraptor "may entirely mislead us as to its feeding habits and belie its character". In the 1990s, the discovery of nesting oviraptorids like Citipati proved that Osborn was correct in his caution regarding the name. These finds showed that the eggs in question probably belonged to Oviraptor itself, and that the specimen was actually brooding its eggs, when it died at the nest.

Oviraptor lived in the late Cretaceous period, during the late Campanian stage about 75 million years ago; only one definitive specimen is known (with associated eggs), from the Djadokhta Formation of Mongolia, though a possible second specimen (also with eggs) comes from the northeast region of Inner Mongolia, China, in an area called Bayan Mandahu.

Oviraptoridae

Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between one and two metres long in most cases, though some possible oviraptorids were enormous. Oviraptorids are currently known only from the Late Cretaceous of Asia, with the most well-known species and complete specimens found only in the Gobi Desert of Mongolia and northwestern China.

Oviraptorosauria

Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or without bony crests atop the head. They ranged in size from Caudipteryx, which was the size of a turkey, to the 8-metre-long, 1.4-ton Gigantoraptor. The group (along with all maniraptoran dinosaurs) is close to the ancestry of birds. Analyses like those of Maryanska et al (2002) and Osmólska et al. (2004) suggest that they may represent primitive flightless birds. The most complete oviraptorosaur specimens have been found in Asia. The North American oviraptorosaur record is sparse.The earliest and most basal ("primitive") known oviraptorosaurs are Ningyuansaurus wangi, Protarchaeopteryx robusta and Incisivosaurus gauthieri, both from the lower Yixian Formation of China, dating to about 125 million years ago during the Aptian age of the early Cretaceous period. A tiny neck vertebra reported from the Wadhurst Clay Formation of England shares some features in common with oviraptorosaurs, and may represent an earlier occurrence of this group (at about 140 million years ago).

Tongtianlong

Tongtianlong (meaning "Tongtianyan dragon") is a genus of oviraptorid theropod dinosaurs that lived in the late Maastrichtian epoch of the late Cretaceous period. It contains one species, T. limosus.

Basal oviraptorosaurs
Caudipteridae
Caenagnathoidea

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