Achillobator

Achillobator (/əˌkɪləˈbaɪtɔːr/ ə-KIL-ə-BAY-tor) is a dromaeosaurid theropod dinosaur that lived roughly 93 to 80 million years ago during the Late Cretaceous in what is now Mongolia, in Asia. It was among the largest dromaeosaurs; the holotype and only known individual of Achillobator is estimated at 5 to 6 m (16.4 to 19.7 ft) long.[1][2] Achillobator was a moderately-built, ground-dwelling, bipedal carnivore. It would have been an active predator, hunting with the enlarged, sickle-shaped claw on the second toe.

Achillobator
Temporal range: Late Cretaceous, 93–80 Ma
Achillobator giganticus skeleton
Skeletal restoration by Jaime Headden
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Dromaeosauridae
Clade: Eudromaeosauria
Subfamily: Dromaeosaurinae
Genus: Achillobator
Perle, Norell, & Clark, 1999
Species:
A. giganticus
Binomial name
Achillobator giganticus
Perle, Norell, & Clark, 1999

Discovery and species

Achillobator scale
Size comparison between Achillobator and a human

Fossils of Achillobator were first discovered during a Mongolian and Russian field expedition, and collected by Burkhant in 1989, but the specimen was not described and named until 1999,[3] by Mongolian paleontologist Altangerel Perle, and American paleontologists Mark Norell and Jim Clark, although the description was not complete and was actually published without the knowledge of the latter two paleontologists.

The fossils of the type specimen of Achillobator, FR.MNUFR 15, were found associated but mostly disarticulated, and include a left maxilla with teeth, two cervical vertebrae, two dorsal vertebrae, rib fragments, seven caudal vertebrae, a scapula and coracoid, a pelvis with a right ilium, pubis and ischium, a radius, an incomplete manus, a left femur and tibia, and an incomplete pes. Smith at al. (2012) noted that this genus represents the second largest of the known dromaeosaurid taxa with a tibial length of 490 mm. Its femur, which is 3% longer than the tibia, a rare trait in Dromaeosaurs, measures 505 mm in length. Estimates suggest that Achillobator weighed 350 kg (771.6 lb) at most.[4] The teeth are serrated and recurved, and the posterior serrations are slightly larger than the anterior serrations.

Achillobator reconstruction
Restoration

The genus name Achillobator means 'Achilles hero' and is derived from Achilles, a famous ancient Greek warrior who fought in the Trojan War, and the Mongolian word baatar, anciently bagatur, which means 'hero'. The generic name refers to the large Achilles tendon that connects to the sickle claw on the foot, which was the major combat weapon of dromaeosaurids. The specific name giganticus, is derived from Greek word gigantas (γίγαντας) meaning 'giant', which is in reference to Achillobator's size, which exceeds that of most other dromaeosaurids.

Chimera hypothesis

The pelvis of Achillobator seems to show plesiomorphic ("primitive") saurischian characteristics compared to other dromaeosaurids. For instance, the pubis is aligned vertically and has a large pubic boot (a wide expansion at the end), unlike most other dromaeosaurids, where there is a much smaller boot, if any, and the pubis points backwards in the same direction as the ischium (a condition called opisthopuby, which is also seen in the unrelated therizinosaurs and ornithischians, as well as in birds).

The above differences and others led Burnham et al. (2000) to suggest that Achillobator represents a paleontological chimera.[5] However, other studies have attempted to refute this, noting that many pieces were found to be semi-articulated, all of the elements are the same color and preservation, and that Achillobator routinely comes out as a dromaeosaurid in cladistic analyses, even taking into account the differences.[6]

Classification

Eudromaeosauria

Bambiraptor

Velociraptorinae

Adasaurus

Tsaagan

Velociraptor

Dromaeosaurinae

Deinonychus

Achillobator

Dromaeosaurus

Utahraptor

Yurgovuchia

Eudromaeosauria cladogram based on the phylogenetic analysis, conducted by Phil Senter, James I. Kirkland, Donald D. DeBlieux, Scott Madsen and Natalie Toth., in 2012.[7]

Achillobator is a dromaeosaurid, a family of dinosaurs currently thought to be very closely related to birds.[8] While the relationship of dromaeosaurids to other theropods (including birds) is relatively well understood, the phylogeny within the family itself is not. The more recent phylogenetic analyses, conducted by Longrich and Currie (2009) and Senter et al. (2012) show that Achillobator is a member of the subfamily Dromaeosaurinae, most closely related to North American forms like Utahraptor and Dromaeosaurus.[9] Deinonychus and Velociraptor are also dromaeosaurids, but appear to represent a different branch of the family.[7][10]

Distinguishing anatomical features

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group.

According to the revised diagnosis by Turner et al. (2012), Achillobator can be distinguished based on the following combination of characteristics and autapomorphies:

  • the promaxillary fenestra is completely exposed
  • the promaxillary and maxillary fenestra are elongate and vertically oriented at same level in the maxilla*
  • metatarsal III is wide proximally
  • the femur is longer than the tibia
  • the pelvis is propubic
  • the obturator process on the ischium is large and triangular, and is situated on the proximal half of ischial shaft
  • the boot at distal symphysis of pubis is both cranially and caudally developed

Paleoecology

Bayan Shireh Formation Fauna
Fauna of Bayan Shireh (Achillobator in light yellow)

The remains of Achillobator were recovered in the Bayan Shireh Formation of Dornogovi Province, Mongolia, in fine-grained, medium sandstone/gray mudstone that was deposited during the Late Cretaceous epoch. The exact age is uncertain, with two competing hypotheses; based on comparisons with other formations, the Bayan Shireh fauna seems to correspond best with the Turonian through early Campanian stages of the Late Cretaceous, about 93 to 80 million years ago.[11] However, examination of the magnetostratigraphy of the formation seems to confirm that the entire Bayan Shireh lies within the Cretaceous Long Normal, which lasted only until the end of the Santonian stage, giving a possible Cenomanian through Santonian age, or between 98 and 83 million years ago.[12]

Talarurus and Achillobator
Restoration of Achillobator surrounding Talarurus

Contemporaneous paleofauna of the Bayan Shireh Formation, in which, Achillobator was unearthed, included:

See also

References

  1. ^ "Dromaeosaurs". The Theropod Database. Archived from the original on 2012-10-24. Retrieved 2010-02-04.
  2. ^ Holtz, Thomas R. Jr. (2011) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2010 Appendix.
  3. ^ Perle, A., Norell, M.A., and Clark, J. (1999). "A new maniraptoran theropod - Achillobator giganticus (Dromaeosauridae) - from the Upper Cretaceous of Burkhant, Mongolia." Contributions of the Mongolian-American Paleontological Project, 101: 1–105.
  4. ^ "ACHILLOBATOR". DinoChecker.com. Retrieved 24 August 2013.
  5. ^ Burnham, D.A., Derstler, K.L., Currie, P.J., Bakker, R.T., Zhou, Z., and Ostrom, J.H. (2000). "Remarkable new birdlike dinosaur (Theropoda: Maniraptora) from the Upper Cretaceous of Montana." University of Kansas Paleontological Contributions, 13: 1-14.
  6. ^ Norell, M.A. and Makovicky, J.A. (2004). "Dromaeosauridae." In Weishampel, D.B., P. Dodson, and H. Osmolska (eds.). The Dinosauria (2nd Edition). Berkeley: University of California Press. Pp. 196-209.
  7. ^ a b Senter, P., Kirkland, J.I., DeBlieux, D.D., Madsen, S., and Toth, N. (2012). "New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail." PLoS ONE, 7(5): e36790. doi:10.1371/journal.pone.0036790
  8. ^ P. Godefroit, P. J. Currie, H. Li, C. Y. Shang, and Z.-M. Dong. 2008. A new species of Velociraptor (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China. Journal of Vertebrate Paleontology 28(2):432-438
  9. ^ N. R. Longrich and P. J. Currie. 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences
  10. ^ Makovicky, J.A., Apesteguía, S., and Agnolín, F.L. (2005). "The earliest dromaeosaurid theropod from South America." Nature, 437: 1007-1011.
  11. ^ Jerzykiewicz, T. and Russell, D.A. (1991). "Late Mesozoic stratigraphy and vertebrates of the Gobi Basin." Cretaceous Research, 12(4): 345-377.
  12. ^ Hicks, J.F., Brinkman, D.L., Nichols, D.J., and Watabe, M. (1999). "Paleomagnetic and palynological analyses of Albian to Santonian strata at Bayn Shireh, Burkhant, and Khuren Dukh, eastern Gobi Desert, Mongolia." Cretaceous Research, 20(6): 829-850.

External links

Acheroraptor

Acheroraptor is an extinct genus of dromaeosaurid theropod dinosaur known from the latest Maastrichtian Hell Creek Formation of Montana, United States. It contains a single species, Acheroraptor temertyorum. A. temertyorum is one of the two geologically youngest known species of dromaeosaurids, the other being Dakotaraptor, which is also known from Hell Creek.

Altangerel Perle

Altangerel Perle (born 1945) is a Mongolian palaeontologist.

He is employed at the National University of Mongolia. He has described species such as Goyocephale lattimorei, Achillobator giganticus and Erlikosaurus andrewsi. He has been honored by Polish palaeontologist Halszka Osmólska, who named the species Hulsanpes perlei after him.

Austroraptor

Austroraptor ( AW-stroh-RAP-tər) is an extinct genus of dromaeosaurid dinosaur that lived about 70 million years ago during the Cretaceous Period in what is now modern Argentina. Austroraptor was a medium sized, moderately-built, ground-dwelling, bipedal carnivore, that could grow up to 5–6 m (16.4–19.7 ft) long. Its length makes Austroraptor one of the largest dromaeosaurids known, with only Achillobator, Dakotaraptor, and Utahraptor approaching or surpassing it in length. It is the largest dromaeosaur to be discovered in the Southern Hemisphere. Particularly notable about the taxon were its relatively short forearms, much shorter in proportion when compared to the majority of the members of its group.

Bayan Shireh Formation

The Bayan Shireh Formation (also known as Bayanshiree Svita or the Baynshirenskaya Formation) is a geological formation in Burkhant, Mongolia, that dates to the late Cretaceous period. The exact age is uncertain, with two competing hypotheses; based on comparisons with other formations, the Bayan Shireh fauna seems to correspond best with the Turonian through early Campanian stages of the Late Cretaceous, about 93 to 80 million years ago. However, examination of the magnetostratigraphy of the formation seems to confirm that the entire Bayan Shireh lies within the Cretaceous Long Normal, which lasted only until the end of the Santonian stage, giving a possible Cenomanian through Santonian age, or between 98 and 83 million years ago. The fauna of this formation is similar to the nearby Iren Dabasu Formation.

Dakotaraptor

Dakotaraptor is a genus of large carnivorous dromaeosaurid theropod from the Late Cretaceous of North America.

The first fossils of Dakotaraptor were found in South Dakota, United States, in 2005. In 2015, the genus Dakotaraptor received its name, meaning "plunderer of Dakota", when the type species Dakotaraptor steini was described. The fossils contain an incomplete skeleton without a skull and some individual bones. They have been found in the Maastrichtian-age Hell Creek Formation, dated to the very end of the Cretaceous period, making Dakotaraptor one of the last surviving dromaeosaurids.

Dakotaraptor was about 5.5 metres (18 ft) long, which makes it one of the largest dromaeosaurids known. It had long arms with one of the lower arm bones showing quill knobs, demonstrating that it was most likely feathered. It also had long rear legs with a very large sickle claw on the second toe; this claw could be used to kill relatively large plant-eating dinosaurs. It lived in the same time and area as many iconic late Cretaceous dinosaurs such as Ankylosaurus, Triceratops, and Tyrannosaurus.

Dromaeosauridae

Dromaeosauridae is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek δρομεῦς (dromeus) meaning 'runner' and σαῦρος (sauros) meaning 'lizard'. In informal usage they are often called raptors (after Velociraptor), a term popularized by the film Jurassic Park; a few types include the term "raptor" directly in their name and have come to emphasize their bird-like appearance and speculated bird-like behavior.

Dromaeosaurid fossils have been found across the globe in North America, Europe, Africa, Asia, South America and Antarctica, with fossilized teeth giving credence to the possibility that they inhabited Australia as well. They first appeared in the mid-Jurassic Period (late Bathonian stage, about 167 million years ago) and survived until the end of the Cretaceous (Maastrichtian stage, 66 ma), existing until the Cretaceous–Paleogene extinction event. The presence of dromaeosaurids as early as the Middle Jurassic has been suggested by the discovery of isolated fossil teeth, though no dromaeosaurid body fossils have been found from this period.

Dromaeosaurinae

Dromaeosaurinae is a subfamily of Dromaeosauridae. Most dromaeosaurines lived in what is now the United States and Canada, as well as Mongolia, and possibly Denmark as well. Isolated teeth that may belong to African dromaeosaurines have also been discovered in Ethiopia. These teeth date to the Tithonian stage, of the Late Jurassic Period.All North American and Asian dromaeosaurine dinosaurs from the Late Cretaceous were generally small, no more than 1.8 metres (5.9 ft) long, in Dromaeosaurus and Adasaurus. However, among the dromaeosaurines were the largest dromaeosaurs ever; Dakotaraptor was ~5.5 metres (18 ft) long, Achillobator 6 metres (20 ft), and Utahraptor up to ~7 metres (23 ft).

Dromaeosaurus

Dromaeosaurus (, "swift running lizard") is a genus of theropod dinosaur which lived during the Late Cretaceous period (middle late Campanian), sometime between 76.5 and 74.8 million years ago, in the western United States and Alberta, Canada. The type species is Dromaeosaurus albertensis, which was described by William Diller Matthew and Barnum Brown in 1922.

Eudromaeosauria

Eudromaeosauria ("true dromaeosaurs") is a subgroup of terrestrial dromaeosaurid theropod dinosaurs. They were relatively large-bodied, feathered hypercarnivores (with diets consisting almost entirely of other terrestrial vertebrates) that flourished in the Cretaceous Period.

Eudromaeosaur fossils are known almost exclusively from the northern hemisphere. They first appeared in the early Cretaceous Period (early Aptian stage, about 124 million years ago) and survived until the end of the Cretaceous (Maastrichtian stage, 66 Ma). The earliest known definitive eudromaeosaur is the dromaeosaurine Utahraptor ostrommaysorum, from the Cedar Mountain Formation, dated to 124 million years ago. However, the earlier (143-million-year-old) fossils such as those of Nuthetes destructor and several indeterminate teeth dating to the Kimmeridgian stage may represent eudromaeosaurs.

Hulsanpes

Hulsanpes is a genus of dromaeosaurid theropod dinosaur from Mongolia that lived during the Late Cretaceous.

Itemirus

Itemirus is a genus of theropod dinosaur from the Turonian age of the Late Cretaceous period of Uzbekistan.

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.

Santonian

The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous epoch or Upper Cretaceous series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The Santonian is preceded by the Coniacian and is followed by the Campanian.

Saurornitholestinae

Saurornitholestinae is a subfamily of dromaeosaurid dinosaurs. The saurornitholestines currently include three monotypic genera: Atrociraptor marshalli, Bambiraptor feinbergorum, and Saurornitholestes langstoni. All are medium-sized dromaeosaurs from the Late Cretaceous of western North America. The group was originally recognized by Longrich and Currie as the sister taxon to a clade formed by the Dromaeosaurinae and Velociraptorinae. However, not all phylogenetic analyses recover this group.

Unenlagiinae

Unenlagiinae is a subfamily of dromaeosaurid theropods. Unenlagiines are known from South America and Antarctica.

Utahraptor

Utahraptor (meaning "Utah's predator" or "Utah's thief") is a genus of theropod dinosaurs. It contains a single species, Utahraptor ostrommaysi, which is the largest-known member of the family Dromaeosauridae. Fossil specimens date to the upper Barremian stage of the early Cretaceous period (in rock strata dated to 126 ± 2.5 million years ago).In 2018, it was proposed that Utahraptor be the Utah state dinosaur, an act that was approved by the Senate. Initially Utahraptor would have replaced another dinosaur, Allosaurus, as the state's official fossil, but it was decided that Utahraptor would be another symbol of the state.

Velociraptor

Velociraptor (; meaning "swift seizer" in Latin) is a genus of dromaeosaurid theropod dinosaur that lived approximately 75 to 71 million years ago during the latter part of the Cretaceous Period. Two species are currently recognized, although others have been assigned in the past. The type species is V. mongoliensis; fossils of this species have been discovered in Mongolia. A second species, V. osmolskae, was named in 2008 for skull material from Inner Mongolia, China.

Smaller than other dromaeosaurids like Deinonychus and Achillobator, Velociraptor nevertheless shared many of the same anatomical features. It was a bipedal, feathered carnivore with a long tail and an enlarged sickle-shaped claw on each hindfoot, which is thought to have been used to tackle and disembowel prey. Velociraptor can be distinguished from other dromaeosaurids by its long and low skull, with an upturned snout.

Velociraptor (commonly shortened to "raptor") is one of the dinosaur genera most familiar to the general public due to its prominent role in the Jurassic Park motion picture series. In real life, however, Velociraptor was roughly the size of a turkey, considerably smaller than the approximately 2 m (7 ft) tall and 80 kg (180 lb) reptiles seen in the films (which were based on members of the related genus Deinonychus). Today, Velociraptor is well known to paleontologists, with over a dozen described fossil skeletons, the most of any dromaeosaurid. One particularly famous specimen preserves a Velociraptor locked in combat with a Protoceratops.

Velociraptorinae

Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. The earliest velociraptorines are probably Nuthetes from the United Kingdom, and possibly Deinonychus from North America. However, several indeterminate velociraptorines have also been discovered, dating to the Kimmeridgian stage, in the Late Jurassic Period. These fossils were discovered in the Langenberg quarry, Oker near Goslar, Germany.In 2007 paleontologists studied front limb bones of Velociraptor and discovered small bumps on the surface, known as quill knobs. The same feature is present in some bird bones, and represents the attachment point for strong secondary wing feathers. This finding provided the first direct evidence that velociraptorines, like all other maniraptorans, had feathers.While most velociraptorines were generally small animals, at least one species may have achieved gigantic sizes comparable to those found among the dromaeosaurines. So far, this unnamed giant velociraptorine is known only from isolated teeth found on the Isle of Wight, England. The teeth belong to an animal the size of dromaeosaurines of the genus Utahraptor, but they appear to belong to a velociraptorine, judging by the shape of the teeth and the anatomy of their serrations.

Yurgovuchia

Yurgovuchia is an extinct genus of dromaeosaurid theropod dinosaur known from the Early Cretaceous (probably Barremian stage) of Utah. It contains a single species, Yurgovuchia doellingi. According to a phylogenetic analysis performed by its describers, it represents an advanced dromaeosaurine, closely related to Achillobator, Dromaeosaurus and Utahraptor.

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