Therizinosaur

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Therizinosauria
Temporal range: EarlyLate Cretaceous, 130–66Ma
Possible Early Jurassic record
Nothronychus (1).jpg
Reconstructed skeleton of Nothronychus mckinleyi
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Suborder: Theropoda
Clade: Maniraptora
Clade: Therizinosauria
Russell, 1997
Synonyms

Segnosauria Barsbold, 1980
Segnosaurischia Dong, 1987

Therizinosaurs (or segnosaurs) are theropod dinosaurs belonging to the clade Therizinosauria. Therizinosaur fossils have been found in Early through Late Cretaceous deposits in Mongolia, the People's Republic of China and Western North America. Various features of the forelimbs, skull and pelvis unite them quite comfortably, both as theropods and, as maniraptorans, close relatives to birds.

The name therizinosaur is derived from the Greek therizo meaning 'to reap' or 'to cut off' and sauros meaning 'lizard'. The older name segnosaur is derived from Latin segnis meaning 'slow' or 'sluggish' and Greek sauros meaning 'lizard'.

Description[edit]

Artist's restoration of Nothronychus mckinleyi

Therizinosaurs had a very distinctive, often confusing set of characteristics. Their long necks, wide torsos, and hind feet with four toes used in walking resembled prosauropod dinosaurs. Their unique hip bones, which pointed backwards and were partially fused together, initially reminded paleontologists of the "bird-hipped" ornithischians. Among the most striking characteristics of therizinosaurs are the enormous claws on their hands, which reached lengths of three feet in Therizinosaurus. The unusual range of motion in therizinosaur forelimbs, which allowed them to reach forward to a degree other theropods could not achieve, also supports the idea that they were mainly herbivorous. Therizinosaurs may have used their long reach and strongly curved claws to grasp and shear leafy branches, in a manner similar to the prehistoric ground sloths.[1]

Skin impressions from Beipiaosaurus indicate that therizinosaurs were covered with a coat of primitive, down-like feathers similar to those seen in the compsognathid Sinosauropteryx, as well as longer, simpler, quill-like feathers that may have been used in display.[2][3] Therizinosaurs spanned a large range of sizes, from the small Beipiaosaurus (which measured 2.2 m, or 7.3 ft in length), to the gigantic Therizinosaurus, which at an approximate 10–12 m (33–40 ft) long and an estimated weight of 6.2 tonnes, was among the largest known theropods.

History[edit]

Outdated life reconstruction of a quadrupedal and prosauropod-like Erlikosaurus andrewsi
Life restoration of Beipiaosaurus inexpectus

Because early finds were incomplete, the strange suite of anatomical features combing features typical of theropods, prosauropods and ornithischians led some scientists, such as Gregory S. Paul, to conclude that segnosaurs (as they were called before Therizinosaurus was recognized as part of the group) represented a late-surviving suborder of primitive dinosaurs, sometimes thought of as intermediates between prosauropods and ornithischians. Because of their suspected relationship with prosauropods, early depictions of segnosaurs (including illustrations by Paul) portrayed them as semi-quadrupedal, a mode of locomotion now known to have been impossible given the bird-like nature of their wrists.[4] It also led Paul to include segnosaurs within paleontologist Robert T. Bakker's Phytodinosauria in 1986, a superorder which was to include ornithischians, prosauropods, and sauropods, typified by their "blunt, spoon-crowned teeth suitable for cropping plants."[4]

It was not until the mid-1990s, after Alxasaurus was discovered and shown to possess more typically theropod features, and Therizinosaurus was recognized as a member of the segnosaur group, that their true identity as herbivorous descendants of the carnivorous theropods became generally accepted.[5] The relation between the more derived therizinosaurids and other theropods was greatly elucidated by the discovery of primitive members of the group, such as Beipiaosaurus in 1999 and Falcarius in 2005.[2] The scientists who described Falcarius noted that it seemed to represent an intermediate stage between carnivorous and herbivorous theropods, a sort of "missing link" between predatory maniraptorans and plant-eating therizinosaurs.[6] Although they are now classified as theropods, therizinosaurs had skulls similar to those of sauropods and the shape of their teeth and jaws make it likely that they were herbivores. 21st century in paleontology 20th century in paleontology 19th century in paleontology 2090s in paleontology 2080s in paleontology 2070s in paleontology 2060s in paleontology 2050s in paleontology 2040s in paleontology 2030s in paleontology 2020s in paleontology 2010s in paleontology 2000s in paleontology 1990s in paleontology 1980s in paleontology 1970s in paleontology 1960s in paleontology 1950s in paleontology 1940s in paleontology 1930s in paleontology 1920s in paleontology 1910s in paleontology 1900s in paleontology 1890s in paleontology 1880s in paleontology 1870s in paleontology 1860s in paleontology 1850s in paleontology 1840s in paleontology 1830s in paleontology 1820s in paleontology Nothronychus Therizinosaurus Enigmosaurus Suzhousaurus Erlikosaurus Segnosaurus Neimongosaurus Nanshiungosaurus Erliansaurus Alxasaurus Beipiaosaurus Jianchangosaurus Falcarius 21st century in paleontology 20th century in paleontology 19th century in paleontology 2090s in paleontology 2080s in paleontology 2070s in paleontology 2060s in paleontology 2050s in paleontology 2040s in paleontology 2030s in paleontology 2020s in paleontology 2010s in paleontology 2000s in paleontology 1990s in paleontology 1980s in paleontology 1970s in paleontology 1960s in paleontology 1950s in paleontology 1940s in paleontology 1930s in paleontology 1920s in paleontology 1910s in paleontology 1900s in paleontology 1890s in paleontology 1880s in paleontology 1870s in paleontology 1860s in paleontology 1850s in paleontology 1840s in paleontology 1830s in paleontology 1820s in paleontology

Systematics[edit]

Taxonomy[edit]

Diagram of various skulls

Barsbold and Perle named the group Segnosauria as an infraorder of Theropoda in 1980.[7] Dong Zhiming (1992) went further, placing the segnosaurs in their own order, Segnosaurischia. This name has been abandoned since the discovery that segnosaurs are a specialized group within the suborder Theropoda. Clark et al. in 2004 considered Segnosaurischia a synonym of Therizinosauroidea.

The superfamily Therizinosauroidea had been established by Maleev in 1954, to include only the bizarre, giant-clawed theropod Therizinosaurus. When it was later realized that Therizinosaurus was an advanced segnosaur, Therizinosauroidea was given a phylogenetic definition to include both groups, and has largely replaced the use of the older name Segnosauria in phylogenetic studies, mainly because of the association of the name Segnosauria with the discredited idea that these animals were relatives of prosauropods.

The following taxonomy follows Zanno, 2010 unless otherwise noted.[8]

Phylogeny[edit]

Hips from different genera.

The clade Therizinosauria was first defined by Dale Russell in 1997 as Alxasaurus, Enigmosaurus, Erlikosaurus, Nanshiungosaurus, Segnosaurus, Therizinosaurus, and all taxa closer to them than to oviraptorosaurs, ornithomimids, and troodontids. Paul Sereno, in 2005, modified this definition to the most inclusive clade containing Therizinosaurus but not Ornithomimus, Oviraptor, Shuvuuia, Tyrannosaurus, or Troodon.[9]

Therizinosauroidea, previously named as a superfamily with no phylogenetic definition, was first defined by Zhang et al. in 2001, as the clade containing all theropods more closely related to Therizinosaurus than to birds (effectively replacing the older name Segnosauria, which has not yet been defined as a clade). This definition, however, defines the same group as the pre-existing Therizinosauria. An alternate definition was given by Clark in 2004 (as the last common ancestor of Therizinosaurus and Beipiaosaurus and all its descendants), comprising a narrower group that excludes more primitive therizinosaurs, such as Falcarius, and allows the name Therizinosauria to remain in use for the larger group comprising all therizinosaurs. This definition was followed by Maryanska and Barsbold (2004), Sereno (2005), Zanno et al. (2009) and Zanno (2010),[8][9][10][11] though other subsequent studies, such as Senter (2007, 2012) have continued to use Therizinosauroidea for the therizinosaur "total group".[12]

The following cladogram follows an analysis by Phil Senter, 2007.[12]

Therizinosauroidea

Falcarius


unnamed

Beipiaosaurus


unnamed

Alxasaurus


unnamed

Nanshiungosaurus


Therizinosauridae

Erliansaurus



Nothronychus


unnamed

Neimongosaurus


unnamed

Segnosaurus


unnamed

Erlikosaurus



Therizinosaurus










The cladogram below follows the extensive phylogenetic analysis of Therizinosauria, by Lindsay E. Zanno, 2010.[8]

Therizinosauria

Falcarius


Therizinosauroidea

Beipiaosaurus


unnamed

Alxasaurus


unnamed

Erliansaurus


unnamed

Neimongosaurus


unnamed

Enigmosaurus


unnamed

Suzhousaurus


Therizinosauridae

Nanshiungosaurus



Segnosaurus



Erlikosaurus



Therizinosaurus


Nothronychus

Nothronychus mckinleyi



Nothronychus graffami











The following cladogram is based on the phylogenetic analysis by Phil Senter et al., 2012.[13]

Therizinosauroidea

Falcarius




Beipiaosaurus




Martharaptor




Alxasaurus


Therizinosauridae

Nanshiungosaurus




Suzhousaurus




Nothronychus




Segnosaurus




Neimongosaurus



Erliansaurus




Erlikosaurus



Therizinosaurus












The cladogram below is the most recent cladogram based on the phylogenetic analysis of Therizinosauria conducted by Hanyong Pu et al., 2013.[14]

Therizinosauria

Falcarius


unnamed

Jianchangosaurus


Therizinosauroidea

Beipiaosaurus


unnamed

Alxasaurus


Therizinosauridae

Erliansaurus



Nanshiungosaurus



Neimongosaurus




Segnosaurus



Erlikosaurus



Suzhousaurus



Enigmosaurus



Therizinosaurus



Nothronychus mckinleyi



Nothronychus graffami








Paleobiology[edit]

Therizinosauroid behavior is quite poorly understood, but recent studies and subsequent finds have revealed some aspects of their behavior. Nests have been discovered with sub-spherical eggs found within them. Evidence points to the eggs being buried and abandoned by the parents, and studies of neonates reveal they were well developed and likely precocial, able to leave the nest shortly after birth, suggesting little to no parental care.[15]

CT scans published in 2012 by Stephan Lautenschlager et al., focused on the skull and brain cavity of Erlikosaurus, revealing it to have a large forebrain. Further findings revealed it had well developed senses of balance, hearing and smell, which may have been useful in evading predators, finding food, or in complex social behavior.[16]

In 2011, a nesting ground consisting of 17 clutches of eggs were found in Mongolia's Gobi Desert, with a total of 75 eggs having been uncovered during excavation. The presence of so many fossilized eggs showed that Therizinosaurs probably were social animals, and that they may have come together for nesting and that some genera may have had parental care. The eggs were 5 inches in diameter and contained no embryos, with evidence indicating the young had hatched and left with their parents in the form of egg shells that had been broken out of. The adult therizinosaurs were estimated to have weighed around 220 pounds from estimates made. The find was described by Yoshitsugu Kobayashi et al. The find, along with mass-death quarries such as those belonging to Falcarius, also further cements evidence that Therizinosauroids were social, herding animals. [17][18]

References[edit]

  1. ^ * Burch, S. (2006). "The range of motion of the glenohumeral joint of the therizinosaur Neimongosaurus yangi (Dinosauria: Theropoda)." Chicago Biological Investigator, 3(2): 20. (Abstract).
  2. ^ a b Xu, X.; Tang, Z-L.; Wang, X-L. (1999). "A therizinosauroid dinosaur with integumentary structures from China". Nature 399 (6734): 350–354. doi:10.1038/20670. 
  3. ^ Xu X., Zheng X.-t. and You, H.-l. (2009). "A new feather type in a nonavian theropod and the early evolution of feathers." Proceedings of the National Academy of Sciences (Philadelphia), . doi:10.1073/pnas.0810055106
  4. ^ a b Paul, G.S. (1988). Predatory Dinosaurs of the World, a Complete Illustrated Guide. New York: Simon and Schuster. 464 p.
  5. ^ Russell, D.A.; Dong, Z. (1993). "The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People's Republic of China." In Currie, P.J. (ed.).". Results from the Sino-Canadian Dinosaur Project. Canadian Journal of Earth Sciences 30: 2107–2127. 
  6. ^ Kirkland, J.I.; Zanno, L.E.; Sampson, S.D.; Clark, J.M.; DeBlieux, D.D. (2005). "A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah". Nature 435 (7038): 84–87. doi:10.1038/nature03468. PMID 15875020. 
  7. ^ Barsbold, R.; Perle, A. (1980). "Segnosauria, a new infraorder of carnivorous dinosaurs". Acta Palaeontologica Polonica 25 (2): 187–195. 
  8. ^ a b c Lindsay E. Zanno (2010). "A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)". Journal of Systematic Palaeontology 8 (4): 503–543. doi:10.1080/14772019.2010.488045. 
  9. ^ a b Sereno, P. C. 2005. Stem Archosauria—TaxonSearch [version 1.0, 2005 November 7]
  10. ^ Clark, J.M., Maryanska, T., and Barsbold, R. (2004). "Therizinosauroidea." Pp. 151– 164 in Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria, Second Edition. University of California Press., 861 pp.
  11. ^ Zanno, L.E., Gillette, D.D., Albright, L.B., and Titus, A.L. (2009). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution." Proceedings of the Royal Society B, Published online before print July 15, 2009, doi:10.1098/rspb.2009.1029.
  12. ^ a b Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda)." Journal of Systematic Palaeontology, 5: 429-463 (doi:10.1017/S1477201907002143).
  13. ^ Senter, P.; Kirkland, J. I.; Deblieux, D. D. (2012). "Martharaptor greenriverensis, a New Theropod Dinosaur from the Lower Cretaceous of Utah". In Dodson, Peter. PLoS ONE 7 (8): e43911. doi:10.1371/journal.pone.0043911. PMC 3430620. PMID 22952806.  edit
  14. ^ Pu, H.; Kobayashi, Y.; Lü, J.; Xu, L.; Wu, Y.; Chang, H.; Zhang, J.; Jia, S. (2013). "An Unusual Basal Therizinosaur Dinosaur with an Ornithischian Dental Arrangement from Northeastern China". In Claessens, Leon. PLoS ONE 8 (5): e63423. doi:10.1371/journal.pone.0063423.  edit
  15. ^ Paul, G.S., 2010, "The Princeton Field Guide to Dinosaurs", Princeton University Press p. 157
  16. ^ http://www.sciencedaily.com/releases/2012/12/121219174154.htm
  17. ^ http://news.yahoo.com/nests-big-clawed-dinosaurs-found-mongolia-123729403.html
  18. ^ http://www.livescience.com/40904-therizinosaur-nesting-colony-discovered.html

External links[edit]