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Theropoda (5)* Clades of Medium to Large Herbivores and Carnivores Later in Dinosaur History, These New Coelophysoidea (10) Chasmosaurus Mariscalensis

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Theropoda (5)* Clades of Medium to Large Herbivores and Carnivores Later in Dinosaur History, These New Coelophysoidea (10) Chasmosaurus Mariscalensis Dinosaurs and the Cretaceous Terrestrial Revolution Graeme T. Lloyd*1, Katie E. Davis2, Davide Pisani3, James E. Tarver1, Marcello Ruta1, Manabu Sakamoto1, David W. E. Hone4,5, Rachel Jennings1 and Michael J. Benton1 1Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol BS8 1RJ, UK; 2DEEB, FBLS, Graham Kerr Building, University of Glasgow, Glasgow G12 8QP, UK; 3Laboratory of Evolutionary Biology, Department of Biology, The National University of Ireland, Maynooth, Co. Kildaire, Ireland; 4Bayerischen Staatssammlung für Paläontologie und Geologie, Richard Wagner-Strasse 10, München 80333, Germany; 5Institute of Vertebrate Paleontology and Paleoanthropology, Xizhimenwai Daije 142,100044 Beijing, People’s Republic of China; Correspondence: [email protected] Abstract Figure 2 (below). Majority-rule consensus tree of the reduced Baum Figure 4. A summary version of the supertree (figure 2) drawn against Basal Ornithischia (3)* and Ragan MRP matrix. Abbreviated clade names are: Mam. = stratigraphy. The 11 statistically significant diversification shifts Basal Thyreophora (3)* Huayangosaurus taibaii The observed diversity of dinosaurs reached its highest peak during the mid- and Late Mamenchisauridae, Br. = Brachiosauridae, Her. = present in both the entire tree (figure 2) and at least one Stegosauridae (11) Struthiosaurus austraicus Cretaceous, the 50Myr that preceded their extinction, and yet this explosion of dinosaur diversity Herrerasauridae, Compsog. = time-sliced tree are marked with white arrows denoting Ankylosauria (27) Stormbergia dangershoeki may be explained largely by sampling bias. It has long been debated whether dinosaurs were “Compsognathoidea”, Ornithomimo. = the branch leading to the more speciose clade. Heterodontosauridae (2) part of the Cretaceous Terrestrial Revolution (KTR), from 125–80Myr ago, when flowering plants, Ornithomimosauria, Therizino. = Taxa in bold represent the collapsing of a Pachycephalosauria (16) Basal Ceratopsia (18)* herbivorous and social insects, squamates, birds and mammals all underwent a rapid expansion. Therizinosauroidea, Alvar. = ia larger clade or group, the size of which Avaceratops lammersi Although an apparent explosion of dinosaur diversity occurred in the mid-Cretaceous, Alvarezsauridae and ch is indicated in parentheses. An ‘*’ Ceratopsidae (18) is Basal Euornithopoda (17)* coinciding with the emergence of new groups (e.g. neoceratopsians, ankylosaurid ankylosaurs, Troodon. = th ia indicates the collapsing of a Dryosauridae (2) ni tops Camptosaurus dispar hadrosaurids and pachycephalosaurs), results from the first quantitative study of diversification Troodontidae. r era Eu paraphyletic group and a ‘†’ Basal Iguanodontoidea (16)* O C orn Euhadrosauria (24) applied to a new supertree of dinosaurs show that this apparent burst in dinosaurian diversity in ith that of an extant clade Basal Sauropodomorpha (6)* the last 18 Myr of the Cretaceous is a sampling artefact. Indeed, major diversification shifts op (i.e. birds). Prosauropoda (14)* od Basal Sauropoda (18)* occurred largely in the first one-third of the group’s history. Despite the appearance of new a Neosauropoda (60) Basal Theropoda (5)* clades of medium to large herbivores and carnivores later in dinosaur history, these new Coelophysoidea (10) Chasmosaurus mariscalensis Arrhinoceratops brachyops Pachyrhinosaurus canadensis Yandusaurus hongheensis Neoceratosauria (15) originations do not correspond to significant diversification shifts. Instead, the overall geometry Hexinlusaurus multidens "Szechuanoraptor" zigongensis Pentaceratops sternbergii Agilisaurus louderbacki ia Einiosaurus procurvicornisChasmosaurus irvinensis Megalosauroidea (14) of the Cretaceous part of the dinosaur tree does not depart from the null hypothesis of an equal Anchiceratops ornatus r Pachyrhinosaurus n. sp. Chasmosaurus russelli Bagaceratops rozhdestvenskyi u Graciliceratops mongoliensisStyracosaurus albertensis Centrosaurus brinkmani Triceratops horridus Carnosauria (12) Achelousaurus horneri a Diceratops hatcheri rates model of lineage branching. Furthermore, we conclude that dinosaurs did not experience Zuniceratops christopheri s Montanoceratops cerorhynchus Chasmosaurus belli o Protoceratops hellenikorhinus Basal Neotetanura (3)* Centrosaurus apertus l Torosaurus latus Udanoceratops tschizhovi Siamotyrannus isanensis a progressive decline at the end of the Cretaceous, nor was their evolution driven directly by the a Prenoceratops pieganensisProtoceratopsAvaceratops andrewsi lammersi Monoclonius crassus h Yamaceratops dorngobiensis Tyrannosauroidea (15) p ‘Compsognathoidea’ (8) KTR. e LiaoceratopsArchaeoceratops yanzigouensisLeptoceratops oshimai gracilis yc Psittacosaurus mongoliensis Scipionyx samniticus h Basal Coelurosauria (5)* c Psittacosaurus Psittacosaurusguyangensis sinensis Ornithomimosauria (11) a PachycephalosaurusPrenocephale wyomingensis edmontonensisChaoyangsaurus youngi P Xuanhuaceratops niei Basal Maniraptora (2) Sphaerotholus buchholtzae Oviraptorosauria (22) 20 Alvarezsauridae (5) Othnielia rex Orodromeus makelai Zephyrosaurus schaffi Jeholosaurus shangyuanensis Hypsilophodon foxii Gasparinisaura cincosaltensis Prenocephale brevis Parksosaurus warreni Thescelosaurus neglectus Prenocephale goodwini Tenontosaurus dossi Gugyedong maniraptoran Tenontosaurus tilletti Excluded Trees: Included Trees: Muttabarrasaurus langdoni Rhabdodon priscus Zalmoxes robustus Zalmoxes shqiperorum Dryosaurus lettowvorbecki Dryosaurus altus Protarchaeopteryx robusta 18 Camptosaurus dispar Yinlong downsi Lurdusaurus arenatus Uncorroborated trees Independent trees Equijubus normani Stygimoloch spinifer Iguanodon bernissartensis Iguanodon atherfieldensis Eumaniraptora (26)† Ouranosaurus nigeriensis HomalocephaleDwarf calathocercos pachycephalosaur Fukuisaurus tetoriensis Redundant trees Dependent trees Tylocephale gilmorei Altirhinus kurzanovi Nanyangosaurus zhugeii Prenocephale prenes Jinzhousaurus yangi 16 Probactrosaurus gobiensis Eolambia caroljonesa Protohadros byrdi Bactrosaurus johnsoni 1 2 3 4 5 6 7 8 9 10 11 12 Gilmoreosaurus mongoliensis OrnatotholusStegoceras browni validum Telmatosaurus transsylvanicus 14 WannanosaurusGoyocephale yansiensis lattimorei Fontllonga euhadrosaur Lophorhothon atopus 225 200 175 150 125 100 75 Gryposaurus notabilis Brachylophosaurus canadensis Maiasaura peeblesorum Yaverlandia bitholus Edmontosaurus annectens P Time (Ma) 12 Edmontosaurus regalis Heterodontosaurus tucki Kerberosaurus manakini l StormbergiaAbrictosaurusStenopelix dangershoeki valdensisconsors Prosaurolophus maximus a e Naashoibitosaurus ostromi "Kritosaurus" australis t 10 a Nodocephalosaurus kirtlandensis Saurolophus osborni e Pararhabdodon isonensis d Tsintaosaurus spinorhinus o i PinacosaurusAnkylosaurus mephistocephalusSaichania magniventris chulsanensis Jaxartosaurus aralensis r Amurosaurus riabinini s 8 EuoplocephalusTarchia gigantea tutus Charonosaurus jiayinensis a u Parasaurolpohus walkeri u N Source Trees a Lambeosaurus lambei Lambeosaurus magnicristatus r 6 s Corythosaurus casuarius i o TalarurusPinacosaurus plicatospineus grangeri Olorotitan arharensis a Dinosaur diversification: diversification shifts l Tsagantegia longicranialis Hypacrosaurus stebingeri Hypacrosaurus altispinus 4 y Tianzhenosaurus youngi Nipponosaurus sachalinensis k Saturnalia tupiniquim Thecodontosaurus antiquus n Shamosaurus scutatus Pantydraco caducus An alternative approach to analysing diversification is to compare 2 Gobisaurus domoculus Mussaurus patagonicus A Jingshanosaurus xinwaensis Efraasia minor observed tree shape with a null model. The simplest null model is Ammosaurus major 0 Polacanthus foxii Riojasaurus incertus equal-rates Markov (ERM), where all branches have the same GargoyleosaurusMymoorapelta parkpinorum maysi Lufengosaurus huenei 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 AnimantarxGastonia ramaljonesi burgei "Gyposaurus" sinensis Minmi paravertebra Massospondylus carinatus probability of splitting resulting in a fully balanced tree. Significant 1985 1987 1989 1991 1993 1995 1997 1999 2001 2003 2005 Yunnanosaurus huangi Edmontonia rugosidens Plateosaurus gracilis Publication Year Plateosaurus cullingworthi excursions from stochastic variation around this model indicates a Edmontonia longiceps Coloradisaurus brevis Panoplosaurus mirus Plateosaurus engelhardti more complex two-parameter model, where, following a split, one Figure 1. Filtering of source trees results in decreased redundancy Unaysaurus tolentinoi SauropeltaSilvisaurus edwardsorum condrayi Anchisaurus polyzelus and a strong skew towards more recent analyses. Uncorroborated Pawpawsaurus campbelli Melanorosaurus readi branch shows a greater probability of splitting than the other. This Cedarpelta bilbeyhallorum Camelotia borealis Lessemsaurus sauropoides M is a diversification shift (Chan and Moore 2005). Here 11 trees lack a numerical cladistic analysis and redundant trees have e Hylaeosaurus armatus Blikanasaurus cromptoni a a Antetonitrus ingenipes m significant shifts (which were also found on time-sliced trees, a been superceded by more recent work. Independent trees StruthiosaurusHungarosaurus austriacus tormai Gongxianosaurus shibeiensis d Isanosaurus attavipachi . i way of testing a potential bias in the assumptions of the ERM r Tazoudasaurus naimi represent a unique combination of taxa and characters, whereas Stegosaurus stenops Kotasaurus yamanpalliensis u LexovisaurusStegosaurus durobrivensis
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