Project 660: Turner, A.; Makovicky, P.; Norell, M.A. 2012. A Review of Dromaeosaurid Systematics and Paravian Phylogeny. Bulletin of the American Museum of Natural History. (371):1-206.
Specimen: †Velociraptor mongoliensis (IGM:100/982)
View: lateral skull

Abstract

Coelurosauria is the most diverse clade of theropod dinosaurs. Much of this diversity is present in Paraves—the clade of dinosaurs containing dromaeosaurids, troodontids, and avialans. Paraves has over 160 million years of evolutionary history that continues to the present day. The clade represents the most diverse living tetrapod group (there are over 9,000 extant species of Aves – a word used here as synonomous with “bird”), and it is at the root of the paravian radiation. When dromaeosaurids, troodontids, and avialans where diverging from one another, that we find the morphology and soft tissue changes associated with the origin of modern avian flight. Within the first 15 million years of known paravian evolutionary history members of this clade exhibited nearly 4 orders of magnitude difference in body size, a value that is similar to the extreme body size disparity present today in mammalian carnivorans, avians, or varanoid squamates. In this respect, Paraves is as an important case study in characterizing the patterns, processes, and dynamics of evolutionary size change. This last point is of particular interest because of the historical significance placed on the role of body size reduction played in the origin of powered avian flight.
Our study reviews and revises the membership of Dromaeosauridae and provides an apomorphy-based diagnosis for all valid taxa. Of the currently 31 named dromaeosaurid species, we found 26 to be valid. We provide the most detailed and comprehensive phylogenetic analysis of paravians to date in order to explore the phylogenetic history of dromaeosaurid taxa. The general pattern of paravian relationships is explored within the broader context of Coelurosauria with an emphasis on sampling basal avialans because of their importance for character optimizations at the base of Paraves.
A large dataset was constructed by merging two datasets, one examining coelurosaur relationships broadly (based on previous TWiG datasets) and the other examining avialan relationships specifically (Clarke et al., 2006). This merged dataset was then significantly revised and supplemented with novel character analysis focusing on paravian taxa. During character analysis, particular attention was given to basal members of Dromaeosauridae, enigmatic basal paravians such as Jinfengopteryx elegans and Anchiornis huxleyi, and the incorporation of new morphological information from two undescribed troodontid species from the Late Cretaceous of Mongolia. A final dataset of 474 characters scored for 111 taxa was used to address paravian evolution. This dataset is important in that it bridges a phylogenetic gap that had persisted between studies on birds and studies on all other coelurosaurs. Most of the scorings in this matrix were based on the direct observation of specimens.
All the most parsimonious trees recovered in the cladistic analysis support the monophyly of Paraves, Troodontidae, Dromaeosauridae, and Deinonychosauria. A new clade of basal troodontids is discovered including two undescribed Mongolian troodontids and Jinfengopteryx elegans. Xiaotingia and Anchiornis form a clade at the base of Troodontidae. Recently proposed relationships within Dromaeosauridae are further supported and a succession of clades from Gondwana and Asia form sister taxa to a clade of Laurasian dromaeosaurids. Avialan monophyly is strongly supported with Archaeopteryx, Sapeornis, Jeholornis, and Jixiangornis forming the successive sister taxa to the Confuciusornis node. This topology supports a more basal position for Sapeornis than previous phylogenetic analyses and indicates a progressive acquisition of a fully “avian” shoulder morphology.


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This project containsMatrices
  • 3 Media
  • 1 Matrix
  • 2 Documents
  • 117 Taxa
  • 3 Specimens
  • 477 Characters
Total size of project's media files: 24 MB
Total scored cells: 24742
Total media associated with cells: 0
Total labels associated with cell media: 0
Characters
Total characters: 477
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 1098
Total character states with associated media: 0
Total character states with media with labels:0
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MorphoBank Project 660


Members

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(scored, NPA, "-")
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Maureen Admin
Project Administrator
Last logged in 01/12/12
117334770024742
(23147, 0, 1595)
000
Peter Makovicky
Full membership
Last logged in Never
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(0, 0, 0)
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Mark Norell
Full membership
Last logged in 01/13/12
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(0, 0, 0)
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Alan Turner
Full membership
Last logged in 08/25/12
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(0, 0, 0)
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Taxonomic Overview for Matrix 'M1507' (117 Taxa)

taxon unscored
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[1] Allosaurus fragilis
Last Modified in 01/12/12
324250022004
[2] Sinraptor dongi
Last Modified in 01/12/12
1393260013004
[3] Ingenia yanshani
Last Modified in 01/12/12
2242270027004
[4] Citipati osmolskae
Last Modified in 01/12/12
464010033004
[5] Rinchenia mongoliensis
Last Modified in 01/12/12
3531040020004
[6] Oviraptor philoceratops
Last Modified in 01/12/12
2991530025004
[7] Conchoraptor gracilis
Last Modified in 01/12/12
2172390021004
[8] Incisivosaurus gauthieri
Last Modified in 01/12/12
313156008004
[9] Microvenator celer
Last Modified in 01/12/12
3111550011004
[10] Chirostenotes pergracilis
Last Modified in 01/12/12
2731810024004
[11] Caudipteryx zoui
Last Modified in 01/12/12
2232290026004
[12] Dromaeosaurus albertensis
Last Modified in 01/12/12
328144005004
[13] Deinonychus antirrhopus
Last Modified in 01/12/12
1053610012004
[14] †Velociraptor mongoliensis
Last Modified in 01/12/12
174470015004
[15] Balaur bondoc
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323149007004
[16] Atrociraptor marshalli
Last Modified in 01/12/12
43939000004
[17] Utahraptor
Last Modified in 01/12/12
371105002004
[18] Adasaurus mongoliensis
Last Modified in 01/12/12
254218007004
[19] Achillobator giganticus
Last Modified in 01/12/12
346127004004
[20] Tsaagan mangas
Last Modified in 01/12/12
276193008004
[21] Saurornitholestes langsto
Last Modified in 01/12/12
349127001004
[22] Bambiraptor feinbergorum
Last Modified in 01/12/12
813880010004
[23] Tianyuraptor ostromi
Last Modified in 01/12/12
332139006004
[24] Sinornithosaurus millenii
Last Modified in 01/12/12
2162510011004
[25] Microraptor zhaoianus
Last Modified in 01/12/12
215258005004
[26] Graciliraptor lujiatunensis
Last Modified in 01/12/12
41163004004
[27] Hesperonychus elizabethae
Last Modified in 01/12/12
46116000004
[28] Pyroraptor olympius
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46116000004
[29] Rahonavis ostromi
Last Modified in 01/12/12
323153002004
[30] Buitreraptor gonzalozorum
Last Modified in 01/12/12
223249009004
[31] Neuquenraptor + unenlagia
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330144003004
[32] Austroraptor
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39780002004
[33] Unenlagia
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362113002004
[34] Unenlagia comahuensis
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369106002004
[35] Unenlagia paynemili
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44631001004
[36] Neuquenraptor argentinus
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44036001004
[37] Shanag ashile
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43936002004
[38] Mahakala omnogovae
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295177008004
[39] Mononykus olecranus
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2661860026004
[40] Shuvuuia deserti
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713750034004
[41] Patagonykus puertai
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3561110012004
[42] Albinykus
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43045002004
[43] Alvarezsaurus calvoi
Last Modified in 01/12/12
40869001004
[44] Proceratosaurus bradleyi
Last Modified in 01/12/12
39580002004
[45] Ornitholestes hermanni
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1842790016004
[46] Coelurus fragilis
Last Modified in 01/12/12
344130003004
[47] Archaeopteryx lithographi
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90384005004
[48] Avimimus portentosus
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2631990016004
[49] Confuciusornis sanctus
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1373320012004
[50] Struthiomimus altus
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1033430032004
[51] Gallimimus bullatus
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553860038004
[52] Garudimimus brevipes
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2542010022004
[53] Pelecanimimus polydon
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374870016004
[54] Harpymimus okladnikovi
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2861700022004
[55] Troodon formosus
Last Modified in 01/12/12
299175003004
[56] Saurornithoides mongolien
Last Modified in 01/12/12
305163009004
[57] Saurornithoides junior
Last Modified in 01/12/12
319153006004
[58] Xixiasaurus
Last Modified in 01/12/12
41163003004
[59] Byronosaurus jaffei
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359112006004
[60] Sinornithoides youngi
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2432160019004
[61] Sinovenator changii
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1613070011004
[62] Mei long
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1413240013004
[63] EK troodontid (igm 100/44
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44433000004
[64] Jinfengopteryx elegans
Last Modified in 01/12/12
363108006004
[65] IGM 100/1126
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2162490013004
[66] IGM 100/1323
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3161520010004
[67] Anchiornis huxleyi
Last Modified in 01/12/12
2072480023004
[68] Xiaotingia
Last Modified in 01/12/12
286191000004
[69] Segnosaurus galbinensis
Last Modified in 01/12/12
365107005004
[70] Erlikosaurus andrewsi
Last Modified in 01/12/12
2781780021004
[71] Alxasaurus elesitaiensis
Last Modified in 01/12/12
3431250010004
[72] Tyrannosaurus rex
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424170019004
[73] Tarbosaurus baatar
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40765005004
[74] Albertosaurus sacrophagus
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122348007004
[75] Gorgosaurus libratus
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654010011004
[76] Daspletosaurus
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95376006004
[77] Eotyrannus lengi
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37299006004
[78] Dilong paradoxus
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1363270015004
[79] Shenzhousaurus orientalis
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2851540040004
[80] Ornithomimus edmonticus
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983410039004
[81] Archaeornithomimus asiati
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2601980021004
[82] Anserimimus planinychus
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38985003004
[83] Huaxiagnathus orientalis
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2462110021004
[84] Sinosauropteryx prima
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2332270018004
[85] Compsognathus longipes
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1952590025004
[86] Juravenator starki
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2282230026004
[87] Jeholornis prima
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1832690025004
[88] Jixiangornis orientalis
Last Modified in 01/12/12
1712810026004
[89] Yanornis martini
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268213002004
[90] Apsaravis ukhaana
Last Modified in 01/12/12
2362260020004
[91] Yixianornis
Last Modified in 01/12/12
1842870011004
[92] Sapeornis
Last Modified in 01/12/12
1612940025004
[93] Neuquenornis volans
Last Modified in 01/12/12
365113002004
[94] Patagopteryx deferrariisi
Last Modified in 01/12/12
320159004004
[95] Cathayornis
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279198007004
[96] Concornis
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349131003004
[97] Gobipteryx
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369102009004
[98] Vorona
Last Modified in 01/12/12
43145001004
[99] Songlingornis
Last Modified in 01/12/12
43147001004
[100] Pengornis houi
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2861840014004
[101] Hesperornis
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1582690050004
[102] Baptornis
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39486000004
[103] Ichthyornis
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1623030018004
[104] Iaceornis marshii
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38393002004
[105] Limenavis patagonica
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44237000004
[106] Crypturellus undulatus
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174030058004
[107] Lithornis
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1433120030004
[108] Gallus gallus
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184070054004
[109] Crax pauxi
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224110044004
[110] Anas platyrhynchus
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323950052004
[111] Chauna torquata
Last Modified in 01/12/12
204140044004
[112] Hongshanornis longicrest
Last Modified in 01/12/12
2751810025004
[113] Liaoningornis longidigitu
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42353001004
[114] Pedopenna
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45620001004
[115] Epidendrosaurus
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40864005004
[116] Epidexipteryx
Last Modified in 01/12/12
3471220010004
[117] Haplocheirus
Last Modified in 01/12/12
2102540013004


Project views

type number of views Individual items viewed (where applicable)
Total project views10396
Project overview1250
Taxon list4038
Media views1874Media search (1210 views); M101475 (249 views); M101471 (202 views); M101477 (213 views);
Matrix views548Matrix landing page (508 views); Matrix (40 views);
Specimen list1339
Views for media list507
Documents list435
Bibliography405




Project downloads

type number of downloads Individual items downloaded (where applicable)
Total downloads from project258
Media downloads99M101471 (32 downloads); M101477 (30 downloads); M101475 (37 downloads);
Project downloads89
Matrix downloads54Matrix (54 downloads);
Document downloads16Character list (10 downloads); Copy of matrix (6 downloads);