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.
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
Total unordered/ordered characters:477/0
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MorphoBank Project 660

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    member name taxa specimens media chars character
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    (scored, NPA, "-")
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    Maureen Admin
    Project Administrator
    Last logged in 01/12/12
    117334770024742
    (23147, 0, 1595)
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    Peter Makovicky
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    Mark Norell
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    Last logged in 01/13/12
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    Alan Turner
    Full membership
    Last logged in 08/25/12
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    000


    Taxonomic Overview for Matrix 'M1507' (117 Taxa)

    taxon unscored
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    no cell
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    "-" cellscell images labels on
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    [1] Allosaurus fragilis
    Last Modified in 12/05/13
    32425423022004
    [2] Sinraptor dongi
    Last Modified in 12/05/13
    139326325013004
    [3] Ingenia yanshani
    Last Modified in 12/05/13
    224227226027004
    [4] Citipati osmolskae
    Last Modified in 12/05/13
    46401398033004
    [5] Rinchenia mongoliensis
    Last Modified in 12/05/13
    353104104020004
    [6] Oviraptor philoceratops
    Last Modified in 12/05/13
    299153153025004
    [7] Conchoraptor gracilis
    Last Modified in 12/05/13
    217239239021004
    [8] Incisivosaurus gauthieri
    Last Modified in 12/05/13
    31315615608004
    [9] Microvenator celer
    Last Modified in 12/05/13
    311155155011004
    [10] Chirostenotes pergracilis
    Last Modified in 12/05/13
    273181180024004
    [11] Caudipteryx zoui
    Last Modified in 12/05/13
    223229228026004
    [12] Dromaeosaurus albertensis
    Last Modified in 12/05/13
    32814414405004
    [13] Deinonychus antirrhopus
    Last Modified in 12/05/13
    105361360012004
    [14] † Velociraptor mongoliensis
    Last Modified in 12/05/13
    17447445015004
    [15] Balaur bondoc
    Last Modified in 12/05/13
    32314914707004
    [16] Atrociraptor marshalli
    Last Modified in 12/05/13
    439393800004
    [17] Utahraptor
    Last Modified in 12/05/13
    37110510402004
    [18] Adasaurus mongoliensis
    Last Modified in 12/05/13
    25421821607004
    [19] Achillobator giganticus
    Last Modified in 12/05/13
    34612712704004
    [20] Tsaagan mangas
    Last Modified in 12/05/13
    27619319308004
    [21] Saurornitholestes langsto
    Last Modified in 12/05/13
    34912712701004
    [22] Bambiraptor feinbergorum
    Last Modified in 12/05/13
    81388386010004
    [23] Tianyuraptor ostromi
    Last Modified in 12/05/13
    33213913906004
    [24] Sinornithosaurus millenii
    Last Modified in 12/05/13
    216251250011004
    [25] Microraptor zhaoianus
    Last Modified in 12/05/13
    21525825705004
    [26] Graciliraptor lujiatunensis
    Last Modified in 12/05/13
    411636204004
    [27] Hesperonychus elizabethae
    Last Modified in 12/05/13
    461161600004
    [28] Pyroraptor olympius
    Last Modified in 12/05/13
    461161600004
    [29] Rahonavis ostromi
    Last Modified in 12/05/13
    32315315202004
    [30] Buitreraptor gonzalozorum
    Last Modified in 12/05/13
    22324924509004
    [31] Neuquenraptor + unenlagia
    Last Modified in 12/05/13
    33014414403004
    [32] Austroraptor
    Last Modified in 12/05/13
    397807802004
    [33] Unenlagia
    Last Modified in 12/05/13
    36211311302004
    [34] Unenlagia comahuensis
    Last Modified in 12/05/13
    36910610602004
    [35] Unenlagia paynemili
    Last Modified in 12/05/13
    446313001004
    [36] Neuquenraptor argentinus
    Last Modified in 12/05/13
    440363601004
    [37] Shanag ashile
    Last Modified in 12/05/13
    439363602004
    [38] Mahakala omnogovae
    Last Modified in 12/05/13
    29517717408004
    [39] Mononykus olecranus
    Last Modified in 12/05/13
    266186185026004
    [40] Shuvuuia deserti
    Last Modified in 12/05/13
    71375372034004
    [41] Patagonykus puertai
    Last Modified in 12/05/13
    356111109012004
    [42] Albinykus
    Last Modified in 12/05/13
    430454502004
    [43] Alvarezsaurus calvoi
    Last Modified in 12/05/13
    408696801004
    [44] Proceratosaurus bradleyi
    Last Modified in 12/05/13
    395808002004
    [45] Ornitholestes hermanni
    Last Modified in 12/05/13
    184279277016004
    [46] Coelurus fragilis
    Last Modified in 12/05/13
    34413013003004
    [47] Archaeopteryx lithographi
    Last Modified in 12/05/13
    9038438205004
    [48] Avimimus portentosus
    Last Modified in 12/05/13
    263199198016004
    [49] Confuciusornis sanctus
    Last Modified in 12/05/13
    137332328012004
    [50] Struthiomimus altus
    Last Modified in 12/05/13
    103343342032004
    [51] Gallimimus bullatus
    Last Modified in 12/05/13
    55386384038004
    [52] Garudimimus brevipes
    Last Modified in 12/05/13
    254201201022004
    [53] Pelecanimimus polydon
    Last Modified in 12/05/13
    3748787016004
    [54] Harpymimus okladnikovi
    Last Modified in 12/05/13
    286170169022004
    [55] Troodon formosus
    Last Modified in 12/05/13
    29917517503004
    [56] Saurornithoides mongolien
    Last Modified in 12/05/13
    30516316309004
    [57] Saurornithoides junior
    Last Modified in 12/05/13
    31915315206004
    [58] Xixiasaurus
    Last Modified in 12/05/13
    411636303004
    [59] Byronosaurus jaffei
    Last Modified in 12/05/13
    35911211206004
    [60] Sinornithoides youngi
    Last Modified in 12/05/13
    243216215019004
    [61] Sinovenator changii
    Last Modified in 12/05/13
    161307305011004
    [62] Mei long
    Last Modified in 12/05/13
    141324323013004
    [63] EK troodontid (igm 100/44
    Last Modified in 12/05/13
    444333300004
    [64] Jinfengopteryx elegans
    Last Modified in 12/05/13
    36310810806004
    [65] IGM 100/1126
    Last Modified in 12/05/13
    216249248013004
    [66] IGM 100/1323
    Last Modified in 12/05/13
    316152151010004
    [67] Anchiornis huxleyi
    Last Modified in 12/05/13
    207248247023004
    [68] Xiaotingia
    Last Modified in 12/05/13
    28619119100004
    [69] Segnosaurus galbinensis
    Last Modified in 12/05/13
    36510710705004
    [70] Erlikosaurus andrewsi
    Last Modified in 12/05/13
    278178178021004
    [71] Alxasaurus elesitaiensis
    Last Modified in 12/05/13
    343125124010004
    [72] Tyrannosaurus rex
    Last Modified in 12/05/13
    42417416019004
    [73] Tarbosaurus baatar
    Last Modified in 12/05/13
    407656505004
    [74] Albertosaurus sacrophagus
    Last Modified in 12/05/13
    12234834807004
    [75] Gorgosaurus libratus
    Last Modified in 12/05/13
    65401401011004
    [76] Daspletosaurus
    Last Modified in 12/05/13
    9537637606004
    [77] Eotyrannus lengi
    Last Modified in 12/05/13
    372999906004
    [78] Dilong paradoxus
    Last Modified in 12/05/13
    136327326015004
    [79] Shenzhousaurus orientalis
    Last Modified in 12/05/13
    285154152040004
    [80] Ornithomimus edmonticus
    Last Modified in 12/05/13
    98341340039004
    [81] Archaeornithomimus asiati
    Last Modified in 12/05/13
    260198196021004
    [82] Anserimimus planinychus
    Last Modified in 12/05/13
    389858503004
    [83] Huaxiagnathus orientalis
    Last Modified in 12/05/13
    246211210021004
    [84] Sinosauropteryx prima
    Last Modified in 12/05/13
    233227226018004
    [85] Compsognathus longipes
    Last Modified in 12/05/13
    195259257025004
    [86] Juravenator starki
    Last Modified in 12/05/13
    228223223026004
    [87] Jeholornis prima
    Last Modified in 12/05/13
    183269269025004
    [88] Jixiangornis orientalis
    Last Modified in 12/05/13
    171281280026004
    [89] Yanornis martini
    Last Modified in 12/05/13
    26821320702004
    [90] Apsaravis ukhaana
    Last Modified in 12/05/13
    236226221020004
    [91] Yixianornis
    Last Modified in 12/05/13
    184287282011004
    [92] Sapeornis
    Last Modified in 12/05/13
    161294291025004
    [93] Neuquenornis volans
    Last Modified in 12/05/13
    36511311002004
    [94] Patagopteryx deferrariisi
    Last Modified in 12/05/13
    32015915304004
    [95] Cathayornis
    Last Modified in 12/05/13
    27919819107004
    [96] Concornis
    Last Modified in 12/05/13
    34913112503004
    [97] Gobipteryx
    Last Modified in 12/05/13
    3691029909004
    [98] Vorona
    Last Modified in 12/05/13
    431454501004
    [99] Songlingornis
    Last Modified in 12/05/13
    431474501004
    [100] Pengornis houi
    Last Modified in 12/05/13
    286184177014004
    [101] Hesperornis
    Last Modified in 12/05/13
    158269269050004
    [102] Baptornis
    Last Modified in 12/05/13
    394868300004
    [103] Ichthyornis
    Last Modified in 12/05/13
    162303297018004
    [104] Iaceornis marshii
    Last Modified in 12/05/13
    383939202004
    [105] Limenavis patagonica
    Last Modified in 12/05/13
    442373500004
    [106] Crypturellus undulatus
    Last Modified in 12/05/13
    17403402058004
    [107] Lithornis
    Last Modified in 12/05/13
    143312304030004
    [108] Gallus gallus
    Last Modified in 12/05/13
    18407405054004
    [109] Crax pauxi
    Last Modified in 12/05/13
    22411411044004
    [110] Anas platyrhynchus
    Last Modified in 12/05/13
    32395393052004
    [111] Chauna torquata
    Last Modified in 12/05/13
    20414413044004
    [112] Hongshanornis longicrest
    Last Modified in 12/05/13
    275181177025004
    [113] Liaoningornis longidigitu
    Last Modified in 12/05/13
    423535301004
    [114] Pedopenna
    Last Modified in 12/05/13
    456202001004
    [115] Epidendrosaurus
    Last Modified in 12/05/13
    408646405004
    [116] Epidexipteryx
    Last Modified in 12/05/13
    347122120010004
    [117] Haplocheirus
    Last Modified in 12/05/13
    210254254013004


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