Project 660: A. Turner, P. Makovicky, M. A. Norell. 2012. A Review of Dromaeosaurid Systematics and Paravian Phylogeny. Bulletin of the American Museum of Natural History. (371):1-206.
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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Project DOI: 10.7934/P660, http://dx.doi.org/10.7934/P660
This project contains | Matrices |
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Download Project SDD File | 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 Total unordered/ordered characters:477/0 |
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MorphoBank Project 660
MorphoBank Project 660
- Creation Date:
22 December 2011 - Publication Date:
04 September 2012 - Project views: 51674
- Media downloads: 39
- Matrix downloads: 154
This research
supported by
Authors' Institutions
- American Museum of Natural History
- Field Museum of Natural History
- Stony Brook University
Members
member name | taxa | specimens | media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
Maureen Admin Project Administrator | 117 | 3 | 3 | 477 | 0 | 0 | 24742 (23147, 0, 1595) | 0 | 0 | 0 | ||||
Peter Makovicky Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Mark Norell Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Alan Turner Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M1507' (117 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] Allosaurus fragilis Last Modified in 12/05/13 | 32 | 425 | 423 | 0 | 22 | 0 | 0 | 4 |
[2] Sinraptor dongi Last Modified in 12/05/13 | 139 | 326 | 325 | 0 | 13 | 0 | 0 | 4 |
[3] Ingenia yanshani Last Modified in 12/05/13 | 224 | 227 | 226 | 0 | 27 | 0 | 0 | 4 |
[4] Citipati osmolskae Last Modified in 12/05/13 | 46 | 401 | 398 | 0 | 33 | 0 | 0 | 4 |
[5] Rinchenia mongoliensis Last Modified in 12/05/13 | 353 | 104 | 104 | 0 | 20 | 0 | 0 | 4 |
[6] Oviraptor philoceratops Last Modified in 12/05/13 | 299 | 153 | 153 | 0 | 25 | 0 | 0 | 4 |
[7] Conchoraptor gracilis Last Modified in 12/05/13 | 217 | 239 | 239 | 0 | 21 | 0 | 0 | 4 |
[8] Incisivosaurus gauthieri Last Modified in 12/05/13 | 313 | 156 | 156 | 0 | 8 | 0 | 0 | 4 |
[9] Microvenator celer Last Modified in 12/05/13 | 311 | 155 | 155 | 0 | 11 | 0 | 0 | 4 |
[10] Chirostenotes pergracilis Last Modified in 12/05/13 | 273 | 181 | 180 | 0 | 24 | 0 | 0 | 4 |
[11] Caudipteryx zoui Last Modified in 12/05/13 | 223 | 229 | 228 | 0 | 26 | 0 | 0 | 4 |
[12] Dromaeosaurus albertensis Last Modified in 12/05/13 | 328 | 144 | 144 | 0 | 5 | 0 | 0 | 4 |
[13] Deinonychus antirrhopus Last Modified in 12/05/13 | 105 | 361 | 360 | 0 | 12 | 0 | 0 | 4 |
[14] † Velociraptor mongoliensis Last Modified in 12/05/13 | 17 | 447 | 445 | 0 | 15 | 0 | 0 | 4 |
[15] Balaur bondoc Last Modified in 12/05/13 | 323 | 149 | 147 | 0 | 7 | 0 | 0 | 4 |
[16] Atrociraptor marshalli Last Modified in 12/05/13 | 439 | 39 | 38 | 0 | 0 | 0 | 0 | 4 |
[17] Utahraptor Last Modified in 12/05/13 | 371 | 105 | 104 | 0 | 2 | 0 | 0 | 4 |
[18] Adasaurus mongoliensis Last Modified in 12/05/13 | 254 | 218 | 216 | 0 | 7 | 0 | 0 | 4 |
[19] Achillobator giganticus Last Modified in 12/05/13 | 346 | 127 | 127 | 0 | 4 | 0 | 0 | 4 |
[20] Tsaagan mangas Last Modified in 12/05/13 | 276 | 193 | 193 | 0 | 8 | 0 | 0 | 4 |
[21] Saurornitholestes langsto Last Modified in 12/05/13 | 349 | 127 | 127 | 0 | 1 | 0 | 0 | 4 |
[22] Bambiraptor feinbergorum Last Modified in 12/05/13 | 81 | 388 | 386 | 0 | 10 | 0 | 0 | 4 |
[23] Tianyuraptor ostromi Last Modified in 12/05/13 | 332 | 139 | 139 | 0 | 6 | 0 | 0 | 4 |
[24] Sinornithosaurus millenii Last Modified in 12/05/13 | 216 | 251 | 250 | 0 | 11 | 0 | 0 | 4 |
[25] Microraptor zhaoianus Last Modified in 12/05/13 | 215 | 258 | 257 | 0 | 5 | 0 | 0 | 4 |
[26] Graciliraptor lujiatunensis Last Modified in 12/05/13 | 411 | 63 | 62 | 0 | 4 | 0 | 0 | 4 |
[27] Hesperonychus elizabethae Last Modified in 12/05/13 | 461 | 16 | 16 | 0 | 0 | 0 | 0 | 4 |
[28] Pyroraptor olympius Last Modified in 12/05/13 | 461 | 16 | 16 | 0 | 0 | 0 | 0 | 4 |
[29] Rahonavis ostromi Last Modified in 12/05/13 | 323 | 153 | 152 | 0 | 2 | 0 | 0 | 4 |
[30] Buitreraptor gonzalozorum Last Modified in 12/05/13 | 223 | 249 | 245 | 0 | 9 | 0 | 0 | 4 |
[31] Neuquenraptor + unenlagia Last Modified in 12/05/13 | 330 | 144 | 144 | 0 | 3 | 0 | 0 | 4 |
[32] Austroraptor Last Modified in 12/05/13 | 397 | 80 | 78 | 0 | 2 | 0 | 0 | 4 |
[33] Unenlagia Last Modified in 12/05/13 | 362 | 113 | 113 | 0 | 2 | 0 | 0 | 4 |
[34] Unenlagia comahuensis Last Modified in 12/05/13 | 369 | 106 | 106 | 0 | 2 | 0 | 0 | 4 |
[35] Unenlagia paynemili Last Modified in 12/05/13 | 446 | 31 | 30 | 0 | 1 | 0 | 0 | 4 |
[36] Neuquenraptor argentinus Last Modified in 12/05/13 | 440 | 36 | 36 | 0 | 1 | 0 | 0 | 4 |
[37] Shanag ashile Last Modified in 12/05/13 | 439 | 36 | 36 | 0 | 2 | 0 | 0 | 4 |
[38] Mahakala omnogovae Last Modified in 12/05/13 | 295 | 177 | 174 | 0 | 8 | 0 | 0 | 4 |
[39] Mononykus olecranus Last Modified in 12/05/13 | 266 | 186 | 185 | 0 | 26 | 0 | 0 | 4 |
[40] Shuvuuia deserti Last Modified in 12/05/13 | 71 | 375 | 372 | 0 | 34 | 0 | 0 | 4 |
[41] Patagonykus puertai Last Modified in 12/05/13 | 356 | 111 | 109 | 0 | 12 | 0 | 0 | 4 |
[42] Albinykus Last Modified in 12/05/13 | 430 | 45 | 45 | 0 | 2 | 0 | 0 | 4 |
[43] Alvarezsaurus calvoi Last Modified in 12/05/13 | 408 | 69 | 68 | 0 | 1 | 0 | 0 | 4 |
[44] Proceratosaurus bradleyi Last Modified in 12/05/13 | 395 | 80 | 80 | 0 | 2 | 0 | 0 | 4 |
[45] Ornitholestes hermanni Last Modified in 12/05/13 | 184 | 279 | 277 | 0 | 16 | 0 | 0 | 4 |
[46] Coelurus fragilis Last Modified in 12/05/13 | 344 | 130 | 130 | 0 | 3 | 0 | 0 | 4 |
[47] Archaeopteryx lithographi Last Modified in 12/05/13 | 90 | 384 | 382 | 0 | 5 | 0 | 0 | 4 |
[48] Avimimus portentosus Last Modified in 12/05/13 | 263 | 199 | 198 | 0 | 16 | 0 | 0 | 4 |
[49] Confuciusornis sanctus Last Modified in 12/05/13 | 137 | 332 | 328 | 0 | 12 | 0 | 0 | 4 |
[50] Struthiomimus altus Last Modified in 12/05/13 | 103 | 343 | 342 | 0 | 32 | 0 | 0 | 4 |
[51] Gallimimus bullatus Last Modified in 12/05/13 | 55 | 386 | 384 | 0 | 38 | 0 | 0 | 4 |
[52] Garudimimus brevipes Last Modified in 12/05/13 | 254 | 201 | 201 | 0 | 22 | 0 | 0 | 4 |
[53] Pelecanimimus polydon Last Modified in 12/05/13 | 374 | 87 | 87 | 0 | 16 | 0 | 0 | 4 |
[54] Harpymimus okladnikovi Last Modified in 12/05/13 | 286 | 170 | 169 | 0 | 22 | 0 | 0 | 4 |
[55] Troodon formosus Last Modified in 12/05/13 | 299 | 175 | 175 | 0 | 3 | 0 | 0 | 4 |
[56] Saurornithoides mongolien Last Modified in 12/05/13 | 305 | 163 | 163 | 0 | 9 | 0 | 0 | 4 |
[57] Saurornithoides junior Last Modified in 12/05/13 | 319 | 153 | 152 | 0 | 6 | 0 | 0 | 4 |
[58] Xixiasaurus Last Modified in 12/05/13 | 411 | 63 | 63 | 0 | 3 | 0 | 0 | 4 |
[59] Byronosaurus jaffei Last Modified in 12/05/13 | 359 | 112 | 112 | 0 | 6 | 0 | 0 | 4 |
[60] Sinornithoides youngi Last Modified in 12/05/13 | 243 | 216 | 215 | 0 | 19 | 0 | 0 | 4 |
[61] Sinovenator changii Last Modified in 12/05/13 | 161 | 307 | 305 | 0 | 11 | 0 | 0 | 4 |
[62] Mei long Last Modified in 12/05/13 | 141 | 324 | 323 | 0 | 13 | 0 | 0 | 4 |
[63] EK troodontid (igm 100/44 Last Modified in 12/05/13 | 444 | 33 | 33 | 0 | 0 | 0 | 0 | 4 |
[64] Jinfengopteryx elegans Last Modified in 12/05/13 | 363 | 108 | 108 | 0 | 6 | 0 | 0 | 4 |
[65] IGM 100/1126 Last Modified in 12/05/13 | 216 | 249 | 248 | 0 | 13 | 0 | 0 | 4 |
[66] IGM 100/1323 Last Modified in 12/05/13 | 316 | 152 | 151 | 0 | 10 | 0 | 0 | 4 |
[67] Anchiornis huxleyi Last Modified in 12/05/13 | 207 | 248 | 247 | 0 | 23 | 0 | 0 | 4 |
[68] Xiaotingia Last Modified in 12/05/13 | 286 | 191 | 191 | 0 | 0 | 0 | 0 | 4 |
[69] Segnosaurus galbinensis Last Modified in 12/05/13 | 365 | 107 | 107 | 0 | 5 | 0 | 0 | 4 |
[70] Erlikosaurus andrewsi Last Modified in 12/05/13 | 278 | 178 | 178 | 0 | 21 | 0 | 0 | 4 |
[71] Alxasaurus elesitaiensis Last Modified in 12/05/13 | 343 | 125 | 124 | 0 | 10 | 0 | 0 | 4 |
[72] Tyrannosaurus rex Last Modified in 12/05/13 | 42 | 417 | 416 | 0 | 19 | 0 | 0 | 4 |
[73] Tarbosaurus baatar Last Modified in 12/05/13 | 407 | 65 | 65 | 0 | 5 | 0 | 0 | 4 |
[74] Albertosaurus sacrophagus Last Modified in 12/05/13 | 122 | 348 | 348 | 0 | 7 | 0 | 0 | 4 |
[75] Gorgosaurus libratus Last Modified in 12/05/13 | 65 | 401 | 401 | 0 | 11 | 0 | 0 | 4 |
[76] Daspletosaurus Last Modified in 12/05/13 | 95 | 376 | 376 | 0 | 6 | 0 | 0 | 4 |
[77] Eotyrannus lengi Last Modified in 12/05/13 | 372 | 99 | 99 | 0 | 6 | 0 | 0 | 4 |
[78] Dilong paradoxus Last Modified in 12/05/13 | 136 | 327 | 326 | 0 | 15 | 0 | 0 | 4 |
[79] Shenzhousaurus orientalis Last Modified in 12/05/13 | 285 | 154 | 152 | 0 | 40 | 0 | 0 | 4 |
[80] Ornithomimus edmonticus Last Modified in 12/05/13 | 98 | 341 | 340 | 0 | 39 | 0 | 0 | 4 |
[81] Archaeornithomimus asiati Last Modified in 12/05/13 | 260 | 198 | 196 | 0 | 21 | 0 | 0 | 4 |
[82] Anserimimus planinychus Last Modified in 12/05/13 | 389 | 85 | 85 | 0 | 3 | 0 | 0 | 4 |
[83] Huaxiagnathus orientalis Last Modified in 12/05/13 | 246 | 211 | 210 | 0 | 21 | 0 | 0 | 4 |
[84] Sinosauropteryx prima Last Modified in 12/05/13 | 233 | 227 | 226 | 0 | 18 | 0 | 0 | 4 |
[85] Compsognathus longipes Last Modified in 12/05/13 | 195 | 259 | 257 | 0 | 25 | 0 | 0 | 4 |
[86] Juravenator starki Last Modified in 12/05/13 | 228 | 223 | 223 | 0 | 26 | 0 | 0 | 4 |
[87] Jeholornis prima Last Modified in 12/05/13 | 183 | 269 | 269 | 0 | 25 | 0 | 0 | 4 |
[88] Jixiangornis orientalis Last Modified in 12/05/13 | 171 | 281 | 280 | 0 | 26 | 0 | 0 | 4 |
[89] Yanornis martini Last Modified in 12/05/13 | 268 | 213 | 207 | 0 | 2 | 0 | 0 | 4 |
[90] Apsaravis ukhaana Last Modified in 12/05/13 | 236 | 226 | 221 | 0 | 20 | 0 | 0 | 4 |
[91] Yixianornis Last Modified in 12/05/13 | 184 | 287 | 282 | 0 | 11 | 0 | 0 | 4 |
[92] Sapeornis Last Modified in 12/05/13 | 161 | 294 | 291 | 0 | 25 | 0 | 0 | 4 |
[93] Neuquenornis volans Last Modified in 12/05/13 | 365 | 113 | 110 | 0 | 2 | 0 | 0 | 4 |
[94] Patagopteryx deferrariisi Last Modified in 12/05/13 | 320 | 159 | 153 | 0 | 4 | 0 | 0 | 4 |
[95] Cathayornis Last Modified in 12/05/13 | 279 | 198 | 191 | 0 | 7 | 0 | 0 | 4 |
[96] Concornis Last Modified in 12/05/13 | 349 | 131 | 125 | 0 | 3 | 0 | 0 | 4 |
[97] Gobipteryx Last Modified in 12/05/13 | 369 | 102 | 99 | 0 | 9 | 0 | 0 | 4 |
[98] Vorona Last Modified in 12/05/13 | 431 | 45 | 45 | 0 | 1 | 0 | 0 | 4 |
[99] Songlingornis Last Modified in 12/05/13 | 431 | 47 | 45 | 0 | 1 | 0 | 0 | 4 |
[100] Pengornis houi Last Modified in 12/05/13 | 286 | 184 | 177 | 0 | 14 | 0 | 0 | 4 |
[101] Hesperornis Last Modified in 12/05/13 | 158 | 269 | 269 | 0 | 50 | 0 | 0 | 4 |
[102] Baptornis Last Modified in 12/05/13 | 394 | 86 | 83 | 0 | 0 | 0 | 0 | 4 |
[103] Ichthyornis Last Modified in 12/05/13 | 162 | 303 | 297 | 0 | 18 | 0 | 0 | 4 |
[104] Iaceornis marshii Last Modified in 12/05/13 | 383 | 93 | 92 | 0 | 2 | 0 | 0 | 4 |
[105] Limenavis patagonica Last Modified in 12/05/13 | 442 | 37 | 35 | 0 | 0 | 0 | 0 | 4 |
[106] Crypturellus undulatus Last Modified in 12/05/13 | 17 | 403 | 402 | 0 | 58 | 0 | 0 | 4 |
[107] Lithornis Last Modified in 12/05/13 | 143 | 312 | 304 | 0 | 30 | 0 | 0 | 4 |
[108] Gallus gallus Last Modified in 12/05/13 | 18 | 407 | 405 | 0 | 54 | 0 | 0 | 4 |
[109] Crax pauxi Last Modified in 12/05/13 | 22 | 411 | 411 | 0 | 44 | 0 | 0 | 4 |
[110] Anas platyrhynchus Last Modified in 12/05/13 | 32 | 395 | 393 | 0 | 52 | 0 | 0 | 4 |
[111] Chauna torquata Last Modified in 12/05/13 | 20 | 414 | 413 | 0 | 44 | 0 | 0 | 4 |
[112] Hongshanornis longicrest Last Modified in 12/05/13 | 275 | 181 | 177 | 0 | 25 | 0 | 0 | 4 |
[113] Liaoningornis longidigitu Last Modified in 12/05/13 | 423 | 53 | 53 | 0 | 1 | 0 | 0 | 4 |
[114] Pedopenna Last Modified in 12/05/13 | 456 | 20 | 20 | 0 | 1 | 0 | 0 | 4 |
[115] Epidendrosaurus Last Modified in 12/05/13 | 408 | 64 | 64 | 0 | 5 | 0 | 0 | 4 |
[116] Epidexipteryx Last Modified in 12/05/13 | 347 | 122 | 120 | 0 | 10 | 0 | 0 | 4 |
[117] Haplocheirus Last Modified in 12/05/13 | 210 | 254 | 254 | 0 | 13 | 0 | 0 | 4 |
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Media views | 8578 | Media search (5856 views); M101477 (868 views); M101475 (908 views); M101471 (946 views); |
Specimen list | 5125 | |
Views for media list | 2042 | |
Matrix views | 3230 | Matrix landing page (2805 views); Matrix (425 views); |
Documents list | 2161 | |
Taxon list | 24318 | |
Bibliography | 1520 |
Project downloads
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 922 | |
Project downloads | 702 | |
Media downloads | 39 | M101477 (11 downloads); M101475 (14 downloads); M101471 (14 downloads); |
Matrix downloads | 154 | Matrix (154 downloads); |
Document downloads | 27 | Character list (14 downloads); Copy of matrix (13 downloads); |