Advanced Theropod Phylogeny
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Archosaur Phylogeny Mark Norell, George Barrowclough, Joel Cracraft (AMNH), Peter Makovicky (TFM), Brad Livezey (CMNH), James Clark (GWU), John Gatesy (UCR), Chris Brochu (UI) Saurischian diversity All evidence agrees that crocodiles are the closest living relatives of birds. ~10000 species Saurischia Ornithischia a ir Dinosauria C d r ho ur it ~30 species ot n ar r si O Archosauria The Problem…. • Although it has been firmly established that birds are ingroup dinosaurs there have been no detailed phylogenetic treatments of the complete group at fine taxonomic levels. • No large combined analysis of molecular and morphological characters has been undertaken • Our experience working on coelurosaurian phylogeny has taught us that character placement is greatly affected by taxon sampling regimes • Fine-grained taxonomic sampling is key for a clear understanding of macroevolutionary events, such as avian origins and the evolution of associated key morphological and behavioral character systems. 4 winWgehde-rder wome aaeroe saatur Fossilized behavior Goals • Combine all relevant morphological characters into a single large matrix • Examine an extensive series of archosaur taxa • Code taxa at the specimen level • Sequence an extensive series of Neoaves and Crocodylia • Develop a Supermatrix database to store all relevant character and specimen data The Supermatrix • A virtual archive of information • Includes both text and images • Web accessible • Platform independent • Strict version control • Entries can be combined into composites • Data can be output into formats for phylogenetic analysis Progress Supermatrix model developed and site under construction Hundreds of saurischian specimens have been scored & photographed A master list of morphological characters is compiled and being updated Ongoing development and expansion of analyses for sub-clades 0 Lagerpeton chanarensis Marasuchus lilloensis 50 1 Saurischia 2 Pseudolagosuchus major 51 6 Pisanosaurus mertii 53 7 Heterodontosaurus tucki 52 55 8 Lesothosaurus diagnosticus 10 Eoraptor lunensis 54 11 Herrerasaurus ischigualastensis 56 Dinosauria 12 Coelophysis bauri 58 13 Liliensternus liliensterni 57 14 Elaphrosaurus bambergi Saurischia 15 Saturnalia tupiniquim 16 Thecodontosaurus caducus 60 59 17 Thecodontosaurus antiquus Sauropodomorpha Efraasi inor 61 18 a m 21 Ruehleai bedheimensis 64 19 Plateosaurus engelhardti 62 63 20 Plateosaurus gracilis 29 'Kholumolumosaurus ellenbergerorum' 65 72 30 Riojasaurus incertus 24 Massospondylus carinatus 69 67 22 Coloradisaurus brevis 25 Lufengosaurus huenei 66 26 Gyposaurus sinensis 68 27 Yunnanosaurus huangi 23 Mussaurus patagonicus 28 Anchisaurus polyzelus 70 31 Jinshanosaurus xinwaensis 71 33 Melanorosaurus readi 34 Antetonitrus ingenipes 73 35 Lessemsaurus sauropoides 74 36 Blikanasaurus cromptoni 37 Vulcanodon karibaensis 75 38 Kotasaurus yamanpalliensi 76 41 Shunosaurus lii Eusauropoda 81 44 Volkheimeria chubutiensis 80 43 Lappare ntosaurus madagasc 79 40 Barapasaurus tagorei 78 39 Isanosaurus attavipachi 77 42 Omeisaurus maoianus 45 ata osa ru ari s 82 P g u s f a i 47 Diplodocus carnegii 83 48 Camarasaurus lentus 84 Neosauropoda 49 Brachiosaurus brancai Allosaurus Sinosauropteryx Tyrannosaurus Tyrannosaurs Albertosaurus Struthiomimus Ornithomimus Anserimimus Ornithomi- Archaeornithomimus Gallimimus Garudimimus mosaurs Harpymimus Shenzhousaurus } Pelecanimimus Ornitholestes Mononykus Shuuvuia Patagonykus Alvarezsaur Alvarezsaurus } Ingenia Oviraptor Citipati Oviraptor Conchoraptor Chirostenotes Oviraptorosaurs Avimimus Microvenator Caudipteryx } Beipaosaurus Segnosaurus Erlikosaurus Alxasaurus Dromaeosaurus Deinonychus Velociraptor Unenlagia Utahraptor Saurornitholestes Dromaeosaurs Achillobator Adasaurus Ukhaa dromaosaur } Sinornithosaurus Microraptor Sinovenator EK Troodontid Troodon Saurornithoides Saurornithoides Troodontids Byronosaurus Sinornithoides Rahonavis } Theropod Working Group Archeopteryx Confuciusornis Aves http://research.amnh.org/users/ } norell/TWGhome.html = Liaoning taxon NEOAVIAN PHYLOGENY Mayr & Clarke, 2003 Modern Bird Phylogenetics: Scope of the Molecular Data Follow a liberal interpretation of generic limits: ~2450 extant genera to be sampled (expect 82%, or 2112, to be represented by tissues S & A had 760 genera in their "Tapestry." We expect 2.6X coverage in this study. Sequencing partitioned equally between Cracraft, Barrowclough, and Baker labs Furnariidae Dendrocolaptidae Formicariidae Rhinocryptidae Thamnophilidae Conopophagidae Phylogeny of the Furnariidae Furnarii (suboscines) 135 of 148 genera represented many new insights into generic relationships Dendrocolaptidae Formicariidae Rhynocryptidae Thamnophilidae new biogeographic insights into origins of South American avifauna 16 Fringillidae (6) Motacilla Passer Ploceus 23 Prunella Irenidae (2) Nectariniinae (2) Arcanator 22 Modulatrix Promerops Turdinae (2) Muscicapa Cinclus 24 Mimus 15 Sturnus Bombycillidae (2) Regulus (2) Certhia 13 Sitta 21 Polioptila Troglodytes Zosterops Garrulax Sylvia Cisticola Pycnonotus 9 Aegithalos Hirundo (2) 25 Alauda 12 Culicicapa 8 Elminia Parus (2) Remiz Petroicidae (6) Picathartidae (2) 17 Corvus (3) Cyanocitta Corvinella Lanius Monarcha (2) Grallina Melampitta (2) Corcorax Struthidea Manucodia (2) Paradisaea Ptiloris Chaetorhynchus Rhipidura Dicrurus (2) Aegithina Dryoscopus Telophorus Batis 11 Prionops Vanga Artamus Cracticus Gymnorhina Strepera Oreoica Pitohui Oriolus (2) Sphecotheres Paramythia Colluricincla Pachycephala (2) Falcunculus 10 Ptilorrhoa Coracina (2) Lalage 20 Pericrocotus 14 Vireonidae (2) Erpornis Daphoenositta Oscines 18 Cnemophilus Loboparadisaea 19 Philesturnus Callaeatidae Melanocharitidae (4) Orthonychidae (2) Pomatostomidae (2) Ephthianura Meliphaga Pardalotus (2) Meliphagoidea Malurus 7 Ailuroedus Ptilonorhynchus Climacteris Cormobates Climacteridae Menura Menuridae Campylorhamphus Furnarius 6 Formicarius Scytalopus Conopophaga 5 Thamnophilus Mionectes New World Tyrannus Pipra 1 Rupicola 2 Schiffornis Tityra "Suboscines" Neodrepanis 4 Philepitta Psarisomus Old World 3 Smithornis Pitta AcanthisittaÑAcanthisittidae Coracias Gallus Phylogeny and diversification of the largest avian radiation F. K. Barker, A. Cibois, P. Schikler, J. Feinstein, and J. Cracraft. 2004. PNAS 101:11040-11045 All "families" except 1 (Hypocoliidae, endemic to Iran and Iraq!) RAG-1 and RAG-2 nuclear genes (4129 bp) The passerines (passeriforms) arose in Australasian part of Gondwana prior to KT Significant incongruence with DNA hybridization "Tapestry" (e.g., Australasian "Corvida" massively paraphyletic) Phodilus Tyto Ninox Athene Aegolius Otus OW Strix Ciccaba Otus NW Otus Af r. Asio Strigiformes Bubo RAG-1 Ketupa Nyctea Phylogeny of the Owls Barn owls are sister to strigines as expected Australasian Ninox is sister to all other strigines 25 of 31 genera Otus is polyphyletic and represented now the separate clades make more sense biogeographically What remains to be done…. • Visit more collections and sequence/score more taxa • Annealing of subclade matrices to create Supermatrix • Upload information onto Supermatrix website • Development of user-friendly interface to facilitate broad use. • Large-scale simultaneous analysis of all the data.