Temporal range: Early–Late Cretaceous, 130–65.5Ma
Possible Early Jurassic record
|Reconstructed skeleton of Nothronychus mckinleyi|
Segnosauria Barsbold, 1980
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'.
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.
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. 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.
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. 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."
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. 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. 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. 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.
Barsbold and Perle named the group Segnosauria as an infraorder of Theropoda in 1980. 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.
- Branch Therizinosauria
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.
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), though other subsequent studies, such as Senter (2007) have continued to use Therizinosauroidea for the therizinosaur "total group".
The cladogram below follows the most recent and extensive phylogenetic analysis of Therizinosauria to date, by Lindsay E. Zanno, 2010.
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.
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.
- * 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).
- 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.
- 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
- Paul, G.S. (1988). Predatory Dinosaurs of the World, a Complete Illustrated Guide. New York: Simon and Schuster. 464 p.
- 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.
- 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.
- Barsbold, R.; Perle, A. (1980). "Segnosauria, a new infraorder of carnivorous dinosaurs". Acta Palaeontologica Polonica 25 (2): 187–195.
- 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.
- Sereno, P. C. 2005. Stem Archosauria—TaxonSearch [version 1.0, 2005 November 7]
- 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.
- 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.
- Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda)." Journal of Systematic Palaeontology, 5: 429-463 (doi:10.1017/S1477201907002143).
- Paul, G.S., 2010, "The Princeton Field Guide to Dinosaurs", Princeton University Press p. 157