Maniraptora

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Maniraptorans Temporal range: Middle Jurassic–Recent, 164–0 Ma PreЄ Є O S D C P T J K Pg N Possible Early Jurassic record
The hands of Deinonychus (left) and Archaeopteryx (right) are typical of maniraptorans.
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Superorder:	Dinosauria
Order:	Saurischia
Suborder:	Theropoda
Node:	Maniraptoriformes
Branch:	Maniraptora Gauthier, 1986
Subgroups
†Alvarezsauroidea Aviremigia †Alvarezsauridae †Euronychodon †Kakuru †Ornitholestinae †Paronychodon †Therizinosauria †Xinjiangovenator †Yaverlandia †Yixianosaurus
Synonyms
Metornithes Perle et al., 1993

Maniraptora ("hand snatchers") is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period (see Eshanosaurus), and are regarded as surviving today as about 10,000 species of living birds.

Characteristics

Microraptor specimen with feather impressions

Maniraptorans are characterized by long arms and three-fingered hands (though reduced or fused in some lineages), as well as a "half-moon shaped" (semi-lunate) bone in the wrist (carpus). Maniraptorans are the only dinosaurs known to have breast bones (ossified sternal plates). In 2004, Tom Holtz and Halszka Osmólska pointed out six other maniraptoran characters relating to specific details of the skeleton. Unlike most other saurischian dinosaurs, which have pubic bones that point forward, several groups of maniraptorans have an ornithischian-like backwards-pointing hip bone. A backward-pointing hip characterizes the therizinosaurs, dromaeosaurids, avialans, and some primitive troodontids. The fact that the backward-pointing hip is present in so many diverse maniraptoran groups has led most scientists to conclude that the "primitive" forward-pointing hip seen in advanced troodontids and oviraptorosaurs is an evolutionary reversal, and that these groups evolved from ancestors with backward-pointing hips.^[1]

Feathers and flight

Modern pennaceous feathers and remiges are known in the advanced maniraptoran group Aviremigia. More primitive maniraptorans, such as therizinosaurs (specifically Beipiaosaurus), preserve a combination of simple downy filaments and unique elongated quills.^[2][3] Simple feathers are known from more primitive coelurosaurs such as Sinosauropteryx prima, and possibly from even more distantly related species such as the ornithischian Tianyulong confuciusi and the flying pterosaurs. Thus it appears as if some form of feathers or down-like integument would have been present in all maniraptorans, at least when they were young.^[4]

Maniraptora is the only dinosaur group known to have included flying members, though how far back in this lineage flight extends is controversial. Powered and/or gliding flight is believed to have been present in some types of dromaeosaurid, such as Rahonavis and Microraptor.^[5] Other groups, like the Oviraptorosauria, are not known to have been capable of flight, but some scientists have suggested that they could be descended from ancestors which flew. Paul has suggested that this might be the case. Paul has gone as far as to propose that Therizinosauria, Alvarezsauroidea, and the non-maniraptoran group Ornithomimosauria descended from flying ancestors as well.^[6]

Diet

Jinfengopteryx elegans specimen with seeds preserved in the stomach region

Scientists have traditionally assumed that maniraptorans were ancestrally hypercarnivorous, that is, that most non-avialan species primarily ate and hunted only other vertebrates. However, a number of discoveries made during the first decade of the 21st century, as well as re-evaluation of older evidence, began to suggest that maniraptorans were a primarily omnivorous group, including a number of sub-groups that ate mainly plants, insects, or other food sources besides meat. Additionally, phylogenetic studies of maniraptoran relationships began to more consistently show that herbivorous or omnivorous groups were spread throughout the Maniraptora, rather than representing a single side-branch as previously thought. This lead scientists such as Lindsay Zanno to conclude that the ancestral maniraptoran must have been omnivorous, giving rise to several purely herbivorous groups (such as the therizinosaurs, primitive oviraptorosaurs, and some avialans) and that, among non-avians, only one group reverted to pure carnivores (the dromaeosaurids). Most other groups fell somewhere in between the two extremes, with alvarezsaurids and some avialans being insectivorous, and with advanced oviraptorosaurs and troodontids being omnivorous.^[7][8][9]

Classification

The Maniraptora was originally named by Jacques Gauthier in 1986, for a branch-based clade defined as all dinosaurs closer to modern birds than to the ornithomimids. Gauthier noted that this group could be easily characterized by their long forelimbs and hands, which he interpreted as adaptations for grasping (hence the name Maniraptora, which means "hand snatchers" in relation to their 'seizing hands'). In 1994, Thomas R. Holtz attempted to define the group based on the characteristics of the hand and wrist alone (an apomorphy-based definition), and included the long, thin fingers, bowed, wing-like forearm bones, and half-moon shaped wrist bone as key characters. Most subsequent studies have not followed this definition, however, preferring the earlier branch-based definition.

The branch-based definition usually includes the major groups Deinonychosauria, Oviraptorosauria, Therizinosauria, and Aves. Other taxa often found to be maniraptorans include the alvarezsaurs, Ornitholestes.^[4] Several taxa have been assigned to the Maniraptora more definitively, though their exact placement within the group remains uncertain. These forms include the scansoriopterygids, Pedopenna, and Yixianosaurus, and the dubious Bradycneme.

The following cladogram follows Zanno et al. 2009, with omitted clade names after the definitions in Gauthier & De Quieroz (2001) and Sereno (2005).^[7][10][11]

Maniraptora	†Ornitholestes unnamed †Therizinosauria unnamed †Alvarezsauridae (=Alvarezsauroidea) Aviremigia †Oviraptorosauria Paraves Avialae Deinonychosauria †Troodontidae †Dromaeosauridae

Alternate interpretations

In 2002, Czerkas and Yuan reported that some maniraptoran traits, such as a long, backwards-pointed, pubis, short ischia, as well as a perforated acetabulum (a hip socket that is a hole) are apparently absent in Scansoriopteryx. The authors considered it to be more primitive than true theropods, and hypothesized that maniraptorans may have branched off from theropods at a very early point, or may even have descended from pre-theropod dinosaurs.^[12] Zhang et al., in describing the closely related or conspecific specimen Epidendrosaurus, did not report any of the primitive traits mentioned by Czerkas and Yuan, but did find that the shoulder blade of Epidendrosaurus appeared primitive. Despite this, they placed Epidendrosaurus firmly within Maniraptora.^[13]

Technical diagnosis

Holtz and Osmólska (2004) diagnosed the clade Maniraptora based on the following characters: reduced or absent olecranon process of the ulna, greater trochanter and cranial trochanter of the femur fused into a trochanteric crest. An elongated, backwards-pointing pubic bone is present in therizinosauroids, dromaeosaurids, avialans, and the basal troodontid Sinovenator, which suggests that the propubic condition in advanced troodontids and oviraptorosaurs is a reversal.^[1] Turner et al. (2007) named seven synapomorphies that diagnose Maniraptora.^[4]

References

1. ^ ^{a b} Holtz, T.R. and Osmólska, H. (2004). "Saurischia." In Weishampel, Dodson and Osmólska (eds.), The Dinosauria, second edition. Berkeley: University of California Press.
2. ^ 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 c} Turner, A.H.; Pol, D., Clarke, J.A., Erickson, G.M., and Norell, M. (2007). "A basal dromaeosaurid and size evolution preceding avian flight" (pdf). Science 317 (5843): 1378–1381. doi:10.1126/science.1144066. PMID 17823350. http://www.sciencemag.org/cgi/reprint/317/5843/1378.pdf. 
5. ^ Chiappe, L.M. (2007). Glorified Dinosaurs: The Origin and Early Evolution of Birds. Sydney: UNSW Press.
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. ^ ^{a b} 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.
8. ^ Longrich, Nicholas R.; and Currie, Philip J. (2009). "Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae". Cretaceous Research 30 (1): 239–252. doi:10.1016/j.cretres.2008.07.005. 
9. ^ Holtz, T.R.; Jr; Brinkman, D.L.; Chandler, C.L. (1998). "Dental morphometrics and a possibly omnivorous feeding habit for the theropod dinosaur Troodon". GAIA 15: 159–166. 
10. ^ Gauthier, J. and de Queiroz, K. (2001). "Feathered dinosaurs,flying dinosaurs, crown dinosaurs,and the name "Aves"". Pp. 7-41 in Gauthier, J. and L.F. Gall (eds.), New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. New Haven: Peabody Museum of Natural History, Yale University. ISBN 0-912532-57-2.
11. ^ Sereno, P. C., McAllister, S., and Brusatte, S. L. (2005). "TaxonSearch: a relational database for suprageneric taxa and phylogenetic definitions." PhyloInformatics, 8: 1-21.[1]
12. ^ Czerkas, S.A., and Yuan, C. (2002). "An arboreal maniraptoran from northeast China." Pp. 63-95 in Czerkas, S.J. (Ed.), Feathered Dinosaurs and the Origin of Flight. The Dinosaur Museum Journal 1. The Dinosaur Museum, Blanding, U.S.A. PDF abridged version
13. ^ Zhang, F., Zhou, Z., Xu, X. & Wang, X. (2002). "A juvenile coelurosaurian theropod from China indicates arboreal habits." Naturwissenschaften, 89(9): 394-398. doi:10.1007 /s00114-002-0353-8.

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