Therizinosaur

Therizinosaurs (or segnosaurs) were 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 these finds as both theropods and as maniraptorans, close relatives to birds.

The name therizinosaur is derived from the Greek θερίζω[1] therízein, meaning 'to reap' or 'to cut off', and σαῦρος[2] saûros meaning 'lizard'. The older name segnosaur is derived from Latin segnis meaning 'slow' or 'sluggish', and Greek σαυρος, sauros, meaning 'lizard'.

Therizinosauria
Temporal range: EarlyLate Cretaceous, 130–70 Ma
Possible Early Jurassic record
Nothronychus (1)
Reconstructed skeleton of Nothronychus graffami
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Suborder: Theropoda
Clade: Aveairfoila
Clade: Therizinosauria
Russell, 1997
Synonyms

Segnosauria Barsbold, 1980
Segnosaurischia Dong, 1987

Description

Beipiao1mmartyniuk
Life restoration of Beipiaosaurus inexpectus.

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 those of basal sauropodomorph 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 around one meter 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 large mammals that lived later on, such as chalicotheres, ground sloths, great apes, and giant pandas.[3]

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.[4][5] Therizinosaurs spanned a large range of sizes, from the small Beipiaosaurus (which measured 2.2 m [7 ft 3 in] in length),[6] to the gigantic Therizinosaurus, which at an approximate 10 m (33 ft) long and an estimated weight of 5 tonnes,[7] was among the largest-known theropods.

History of research

Erlikosaurus
Outdated reconstruction of a quadrupedal and prosauropod-like Erlikosaurus. Therizinosaurs were often depicted this way until more complete specimens were found

Because early finds were incomplete, the strange suite of anatomical features combining 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.[8] 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."[8]

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.[9] 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.[4] 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.[10] 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.

Systematics

Therizinosaur skeletons
Skeletons to scale

Taxonomy

Barsbold and Perle named the group Segnosauria as an infraorder of Theropoda in 1980.[11] 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.[12]

Phylogeny

Therizinosauria hips variation
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.[14]

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 Teresa Maryańska and Barsbold (2004), Sereno (2005), Zanno et al. (2009) and Zanno (2010),[12][14][15][16] though other subsequent studies, such as Senter (2007, 2012) have continued to use Therizinosauroidea for the therizinosaur "total group".[17]

The following cladogram follows an analysis by Phil Senter, 2007.[17]

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.[12]

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.[18]

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.[19]

Therizinosauria

Falcarius

unnamed

Jianchangosaurus

Therizinosauroidea

Beipiaosaurus

unnamed

Alxasaurus

Therizinosauridae

Erliansaurus

Nanshiungosaurus

Neimongosaurus

Segnosaurus

Erlikosaurus

Suzhousaurus

Enigmosaurus

Therizinosaurus

Nothronychus mckinleyi

Nothronychus graffami

Paleobiology

Therizinosauroid behavior is quite poorly understood, but 20th-century studies and subsequent finds have revealed some aspects of their behavior. Nests with sub-spherical eggs have been found, and evidence points to the eggs being buried and abandoned by the parents. And studies of neonates indicate they were well developed and likely precocial—able to leave the nest shortly after birth and suggesting little to no parental care.[20]

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, and suggesting it had well developed senses of balance, hearing and smell, all of which would have been useful in evading predators, finding food, or in performng complex social behavior.[21]

In 2011, a nesting ground containing 17 clutches of eggs was found in Mongolia's Gobi Desert, with a total of 75 eggs uncovered during excavation. The eggs were 5 inches in diameter and contained no embryos; and there was evidence—in the form of egg shells that had been broken out of—that the young had hatched and left, presumably with their parents. The presence of so many fossilized eggs in one venue implies that therizinosaurs probably were social animals that came together for nesting; and that some genera may have performed parental care. This find, described by Yoshitsugu Kobayashi et al., and mass-death quarries such as those containing Falcarius, augments evidence that therizinosauroids were social, herding animals. Adult therizinosaurs are estimated to have weighed around 500 kg (1,100 lb).[22][23]

References

  1. ^ θερίζω. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  2. ^ σαύρα. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  3. ^ * 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).
  4. ^ a b Xu, X.; Tang, Z-L.; Wang, X-L. (1999). "A therizinosauroid dinosaur with integumentary structures from China". Nature. 399 (6734): 350–354. Bibcode:1999Natur.399..350X. doi:10.1038/20670.
  5. ^ 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 PMID 19139401
  6. ^ Xu, X.; Tang, Z-L.; Wang, X-L. (1999). "A therizinosauroid dinosaur with integumentary structures from China". Nature. 399 (6734): 350–354. Bibcode:1999Natur.399..350X. doi:10.1038/20670.
  7. ^ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 160
  8. ^ a b Paul, G.S. (1988). Predatory Dinosaurs of the World, a Complete Illustrated Guide. New York: Simon and Schuster. 464 p.
  9. ^ 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. Bibcode:1993CaJES..30.2107R. doi:10.1139/e93-183.
  10. ^ 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. Bibcode:2005Natur.435...84K. doi:10.1038/nature03468. PMID 15875020.
  11. ^ Barsbold, R.; Perle, A. (1980). "Segnosauria, a new infraorder of carnivorous dinosaurs". Acta Palaeontologica Polonica. 25 (2): 187–195.
  12. ^ 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.
  13. ^ Paul, G. S. (2016). The Princeton field guide to dinosaurs, second edition. Princeton University Press.
  14. ^ a b Sereno, P. C. 2005. Stem Archosauria—TaxonSearch Archived 2009-01-15 at the Wayback Machine. [version 1.0, 2005 November 7]
  15. ^ 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.
  16. ^ 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.
  17. ^ 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).
  18. ^ Senter, P.; Kirkland, J. I.; Deblieux, D. D. (2012). Dodson, Peter, ed. "Martharaptor greenriverensis, a New Theropod Dinosaur from the Lower Cretaceous of Utah". PLoS ONE. 7 (8): e43911. Bibcode:2012PLoSO...743911S. doi:10.1371/journal.pone.0043911. PMC 3430620. PMID 22952806.
  19. ^ Pu, H.; Kobayashi, Y.; Lü, J.; Xu, L.; Wu, Y.; Chang, H.; Zhang, J.; Jia, S. (2013). Claessens, Leon, ed. "An Unusual Basal Therizinosaur Dinosaur with an Ornithischian Dental Arrangement from Northeastern China". PLoS ONE. 8 (5): e63423. Bibcode:2013PLoSO...863423P. doi:10.1371/journal.pone.0063423. PMC 3667168. PMID 23734177.
  20. ^ Paul, G.S. (2010). The Princeton Field Guide to Dinosaurs. Princeton University Press. p. 157.
  21. ^ "Inside the head of a dinosaur: Research reveals new information on the evolution of dinosaur senses". ScienceDaily.
  22. ^ "Nests of Big-Clawed Dinosaurs Found in Mongolia". Yahoo News. 5 November 2013.
  23. ^ "Nests of Big-Clawed Dinosaurs Found in Mongolia". LiveScience.com.

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