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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Project DOI: 10.7934/P660, http://dx.doi.org/10.7934/P660
This project containsMatrices
  • 3 Media
  • 1 Matrix
  • 2 Documents
  • 117 Taxa
  • 3 Specimens
  • 477 Characters
Total size of project's media files: 24 MB

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
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MorphoBank Project 660

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    Members

    member name taxa specimens media chars character
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    (scored, NPA, "-")
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    medialabels
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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
    0000000
    (0, 0, 0)
    000
    Mark Norell
    Full membership
    Last logged in 01/13/12
    0000000
    (0, 0, 0)
    000
    Alan Turner
    Full membership
    Last logged in 08/25/12
    0000000
    (0, 0, 0)
    000


    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
    Last Modified in 01/12/12
    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
    Last Modified in 01/12/12
    46116000004
    [29] Rahonavis ostromi
    Last Modified in 01/12/12
    323153002004
    [30] Buitreraptor gonzalozorum
    Last Modified in 01/12/12
    223249009004
    [31] Neuquenraptor + unenlagia
    Last Modified in 01/12/12
    330144003004
    [32] Austroraptor
    Last Modified in 01/12/12
    39780002004
    [33] Unenlagia
    Last Modified in 01/12/12
    362113002004
    [34] Unenlagia comahuensis
    Last Modified in 01/12/12
    369106002004
    [35] Unenlagia paynemili
    Last Modified in 01/12/12
    44631001004
    [36] Neuquenraptor argentinus
    Last Modified in 01/12/12
    44036001004
    [37] Shanag ashile
    Last Modified in 01/12/12
    43936002004
    [38] Mahakala omnogovae
    Last Modified in 01/12/12
    295177008004
    [39] Mononykus olecranus
    Last Modified in 01/12/12
    2661860026004
    [40] Shuvuuia deserti
    Last Modified in 01/12/12
    713750034004
    [41] Patagonykus puertai
    Last Modified in 01/12/12
    3561110012004
    [42] Albinykus
    Last Modified in 01/12/12
    43045002004
    [43] Alvarezsaurus calvoi
    Last Modified in 01/12/12
    40869001004
    [44] Proceratosaurus bradleyi
    Last Modified in 01/12/12
    39580002004
    [45] Ornitholestes hermanni
    Last Modified in 01/12/12
    1842790016004
    [46] Coelurus fragilis
    Last Modified in 01/12/12
    344130003004
    [47] Archaeopteryx lithographi
    Last Modified in 01/12/12
    90384005004
    [48] Avimimus portentosus
    Last Modified in 01/12/12
    2631990016004
    [49] Confuciusornis sanctus
    Last Modified in 01/12/12
    1373320012004
    [50] Struthiomimus altus
    Last Modified in 01/12/12
    1033430032004
    [51] Gallimimus bullatus
    Last Modified in 01/12/12
    553860038004
    [52] Garudimimus brevipes
    Last Modified in 01/12/12
    2542010022004
    [53] Pelecanimimus polydon
    Last Modified in 01/12/12
    374870016004
    [54] Harpymimus okladnikovi
    Last Modified in 01/12/12
    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
    Last Modified in 01/12/12
    359112006004
    [60] Sinornithoides youngi
    Last Modified in 01/12/12
    2432160019004
    [61] Sinovenator changii
    Last Modified in 01/12/12
    1613070011004
    [62] Mei long
    Last Modified in 01/12/12
    1413240013004
    [63] EK troodontid (igm 100/44
    Last Modified in 01/12/12
    44433000004
    [64] Jinfengopteryx elegans
    Last Modified in 01/12/12
    363108006004
    [65] IGM 100/1126
    Last Modified in 01/12/12
    2162490013004
    [66] IGM 100/1323
    Last Modified in 01/12/12
    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
    Last Modified in 01/12/12
    424170019004
    [73] Tarbosaurus baatar
    Last Modified in 01/12/12
    40765005004
    [74] Albertosaurus sacrophagus
    Last Modified in 01/12/12
    122348007004
    [75] Gorgosaurus libratus
    Last Modified in 01/12/12
    654010011004
    [76] Daspletosaurus
    Last Modified in 01/12/12
    95376006004
    [77] Eotyrannus lengi
    Last Modified in 01/12/12
    37299006004
    [78] Dilong paradoxus
    Last Modified in 01/12/12
    1363270015004
    [79] Shenzhousaurus orientalis
    Last Modified in 01/12/12
    2851540040004
    [80] Ornithomimus edmonticus
    Last Modified in 01/12/12
    983410039004
    [81] Archaeornithomimus asiati
    Last Modified in 01/12/12
    2601980021004
    [82] Anserimimus planinychus
    Last Modified in 01/12/12
    38985003004
    [83] Huaxiagnathus orientalis
    Last Modified in 01/12/12
    2462110021004
    [84] Sinosauropteryx prima
    Last Modified in 01/12/12
    2332270018004
    [85] Compsognathus longipes
    Last Modified in 01/12/12
    1952590025004
    [86] Juravenator starki
    Last Modified in 01/12/12
    2282230026004
    [87] Jeholornis prima
    Last Modified in 01/12/12
    1832690025004
    [88] Jixiangornis orientalis
    Last Modified in 01/12/12
    1712810026004
    [89] Yanornis martini
    Last Modified in 01/12/12
    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
    Last Modified in 01/12/12
    279198007004
    [96] Concornis
    Last Modified in 01/12/12
    349131003004
    [97] Gobipteryx
    Last Modified in 01/12/12
    369102009004
    [98] Vorona
    Last Modified in 01/12/12
    43145001004
    [99] Songlingornis
    Last Modified in 01/12/12
    43147001004
    [100] Pengornis houi
    Last Modified in 01/12/12
    2861840014004
    [101] Hesperornis
    Last Modified in 01/12/12
    1582690050004
    [102] Baptornis
    Last Modified in 01/12/12
    39486000004
    [103] Ichthyornis
    Last Modified in 01/12/12
    1623030018004
    [104] Iaceornis marshii
    Last Modified in 01/12/12
    38393002004
    [105] Limenavis patagonica
    Last Modified in 01/12/12
    44237000004
    [106] Crypturellus undulatus
    Last Modified in 01/12/12
    174030058004
    [107] Lithornis
    Last Modified in 01/12/12
    1433120030004
    [108] Gallus gallus
    Last Modified in 01/12/12
    184070054004
    [109] Crax pauxi
    Last Modified in 01/12/12
    224110044004
    [110] Anas platyrhynchus
    Last Modified in 01/12/12
    323950052004
    [111] Chauna torquata
    Last Modified in 01/12/12
    204140044004
    [112] Hongshanornis longicrest
    Last Modified in 01/12/12
    2751810025004
    [113] Liaoningornis longidigitu
    Last Modified in 01/12/12
    42353001004
    [114] Pedopenna
    Last Modified in 01/12/12
    45620001004
    [115] Epidendrosaurus
    Last Modified in 01/12/12
    40864005004
    [116] Epidexipteryx
    Last Modified in 01/12/12
    3471220010004
    [117] Haplocheirus
    Last Modified in 01/12/12
    2102540013004


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