DOCSLIB.ORG

Sexual Selection and Macroevolution

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Sexual Selection and Macroevolution Morphology and Evolution of the Ceratopsian Skull Andrew Knapp Submitted in partial fulfilment of the requirements of the Degree of Doctor of Philosophy School of Biological and Chemical Sciences Queen Mary University of London September 2019 Appendix A: Required statement of originality for inclusion in research degree theses I, Andrew Knapp, confirm that the research included within this thesis is my own work or that where it has been carried out in collaboration with, or supported by others, that this is duly acknowledged below and my contribution indicated. Previously published material is also acknowledged below. I attest that I have exercised reasonable care to ensure that the work is original, and does not to the best of my knowledge break any UK law, infringe any third party’s copyright or other Intellectual Property Right, or contain any confidential material. I accept that the College has the right to use plagiarism detection software to check the electronic version of the thesis. I confirm that this thesis has not been previously submitted for the award of a degree by this or any other university. The copyright of this thesis rests with the author and no quotation from it or information derived from it may be published without the prior written consent of the author. Signature: Andrew Knapp Date: 24th September 2019 1 Details of collaboration and publications: Publications arising from this thesis: Knapp A, Knell RJ, Farke AA, Loewen MA and Hone DWE (2018). Patterns of divergence in the morphology of ceratopsian dinosaurs: sympatry is not a driver of divergence. Proc. R. Soc. B. 285: 20180312. http://dx.doi.org/10.1098/rspb.2018.0312 Knapp A, Knell RJ and Hone DWE (2019). Modularity in the skull of the ceratopsian dinosaur Protoceratops and morphometric evidence of a socio-sexually selected origin for the frill. (prepared for submission) Collaborations: Andy Farke and Mark Loewen provided the taxonomic character matrix used in Chapter 2. Sandra Álvarez-Carretero performed the branch length estimations used to construct the time-scaled phylogeny in Chapter 4. Additional publications: O’Brien DM, Allen CE, Van Kleeck MJ, Hone DWE, Knell RJ, Knapp A, Christiansen S and Emlen DJ (2018). On the evolution of extreme structures: static scaling and the function of sexually selected signals. Animal Behaviour. 144; 95 – 108. 2 Abstract Socio-sexual selection has long been recognised as an important driver of evolution and is known to be an important influence on the morphology and behaviour of extant organisms. The long term macroevolutionary effects of socio-sexual selection are difficult to observe because they operate over long timescales. Theoretical work has suggested that socio-sexual selection can possibly influence speciation, adaptation and extinction rates, but these predictions remain largely untested. The fossil record provides a potential solution for testing macroevolutionary hypotheses regarding socio-sexual selection over millions of years. Recent studies have suggested that the growth patterns of cranial ornamentation in Ceratopsian dinosaurs resemble the growth patterns predicted for a trait under socio-sexual selection. Identifying the presence of socio-sexual selection in extinct organisms is problematic, but evidence from extant taxa provide important evidence in detecting its effects on morphology. In this project, I use a combination of traditionally defined character states and three-dimensional geometric morphometrics, on a large scale for the first time in a dinosaur clade, to test competing macroevolutionary hypotheses of socio-sexual selection. I first test the hypothesis that morphological diversity in ceratopsian dinosaurs is driven by species recognition by comparing differences in character states between ceratopsian clades. I evaluate morphological evidence for socio-sexual selection in fossil specimens, employing novel techniques to assess growth and morphological variation in the skull of the ceratopsian that is best-represented by fossil specimens, Protoceratops andrewsi. Lastly, I extend the morphological dataset to encompass other ceratopsian taxa and examine modularity, morphological disparity and evolutionary rates across the clade. My results refute the hypothesis that exaggerated ceratopsian traits evolved for species recognition and provide support for predictions that ceratopsian traits associated with socio-sexual selection, namely the frill and horns, formed distinct units that both developed and evolved at comparatively rapid rates. 3 Contents Page Abstract 3 Contents 4 List of tables 7 List of Figures 8 List of Institutional Abbreviations 10 Summary 11 1 Introduction 1.1 Background 1.1.1 Sexual selection 13 1.1.2 Social selection 15 1.1.3 Predicted effects of socio-sexual selection on macroevolution 16 1.2 Theoretical and empirical evidence for sexual selection 1.2.1 Sexual selection in extant organisms 17 1.2.2 Sexual selection in the fossil record 22 1.2.3 Ceratopsian dinosaurs 27 1.3 Methodology 1.3.1 Geometric morphometrics 30 1.3.2 Modularity and evolutionary rates 31 1.3.3 Data collection 33 1.3.4 Limitations of fossil specimens 34 1.4 Aims and objectives 35 1.5 Hypotheses 36 2 Patterns of divergence in the morphology of ceratopsian dinosaurs: sympatry is not a driver of ornament evolution. 2.1 Abstract 37 2.2 Introduction 38 2.3 Methods 41 2.4 Results 47 2.5 Discussion 53 4 3 Ontogeny and intraspecific variation of the ceratopsian dinosaur Protoceratops andrewsi 3.1 Abstract 57 3.2 Introduction 58 3.3 Methods 3.3.1 Experimental design 61 3.3.2 Statistical analysis 63 3.4 Results 3.4.1 Morphological trends in the skull of Protoceratops andrewsi 65 3.4.2 Variance and whole skull allometry 65 3.4.3 Modularity 70 3.4.4 Sexual dimorphism 74 3.4.5 Module allometry 75 3.4.6 Module disparity 77 3.5 Discussion 79 4 Modularity, morphological disparity and evolutionary rates in the ceratopsian skull 4.1 Abstract 84 4.2 Introduction 85 4.3 Methods 4.3.1 Data collection 91 4.3.2 Effects of size on shape 95 4.3.3 Modularity testing 95 4.3.4 Divergence times estimation 96 4.3.5 Phylogenetic factors affecting shape 97 4.4 Results 4.4.1 Modularity 98 4.4.2 Morphological variation 101 4.4.3 Allometry 105 4.4.4 Phylogenetic signal and evolutionary rates 111 4.4.5 Ceratopsid epiossifications 119 4.4.6 Triceratops, Torosaurus and Nedoceratops 123 4.5 Discussion 4.5.1 Overview 124 4.5.2 Modularity 124 4.5.3 Allometry and phylogenetic signal 127 4.5.4 Evolutionary rates and morphological diversity 129 4.5.5 Epiossifications 131 5 4.5.6 Status of Nedoceratops and Torosaurus 132 4.5.7 Conclusions 134 5 Conclusions 5.1 Summary of findings 135 5.2 Implications of study and future research 5.2.1 Overview 137 5.2.2 Modularity and evolution 138 5.2.3 Evolutionary rates and disparity 139 5.2.4 Allometry, Cope’s rule and heterochrony 140 5.2.5 Species recognition 141 5.2.6 A hypothesis for the role of the frill in ceratopsian evolution 142 5.2.7 Limitations of fossil data and possible solutions 143 Acknowledgements 146 References 148 Appendix 171 6 List of tables Page 2.1 Temporal calibrations and geographical locations of ceratopsian species 43 2.2 z-scores of second order polynomial regression parameters 52 3.1 Results of maximum likelihood and covariance ratio modularity analyses 72 3.2 Results of Hartigan’s dip test for Protoceratops data 74 3.3 Results of ANCOVA for Protoceratops skull size regressions 75 4.1 Results of allometry analyses for ceratopsian dataset 107 4.2 Results of PGLS analysis of evolutionary allometry 107 4.3 Results of phylogenetic signal analysis and net evolutionary rate analysis 113 Results of phylogenetic signal analysis, net evolutionary rates and morphological 121 4.4 disparity for ceratopsids 7 List of figures Page 1.1 The effect of differing species turnover on apparent species richness 23 1.2 Changes in skull shape with ontogeny in specimens of Protoceratops andrewsi, 27 illustrating growth of frill. 1.3 Examples of ceratopsian cranial diversity 28 2.1 Simplified phylogeny and examples of ceratopsian skulls 40 2.2 Time-scaled phylogeny of all known ceratopsian species 45 2.3 Pairwise comparison plots for sympatry categories and character classes 49 2.4 Distributions of simulated second-order polynomial model output parameters 50 2.5 Mean difference values of individual characters for each main sympatry category 51 3.1 Illustration of adult P. andrewsi skeleton 59 3.2 Layout of landmarks on P. andrewsi skull digital mesh model 62 3.3 Principal components analysis of 31 complete P. andrewsi skulls 66 3.4 Results of allometry analysis for whole-skull data 67 3.5 Per-landmark variance in the skull of P. andrewsi 69 3.6 Modules identified in the skull of P. andrewsi from EMMLi analysis 71 3.7 Projected shapes at minimum (Min) size and maximum (Max) size of the five 73 modules identified in P. andrewsi using EMMLi 3.8 Regression of module centroid size and CAC against skull centroid size 76 3.9 Relationship between module Procrustes variance and growth rate and 78 Procrustes variance and within-module correlation 3.10 Front view of projected shape at minimum and maximum size of the skull of 80 Protoceratops andrewsi 3.11 Hypothetical examples of allometric trajectories under different forms of sexual 82 dimorphism 4.1 Time-scaled phylogeny of ceratopsian taxa used in morphometric study 88 4.2 Landmark guide for morphometric study 93 8 4.3 Results of EMMLi modularity test 100 4.4 Phylomorphospace of whole-skull shape data 103 4.5 Phylomorphospaces of individual modules
Load more

Recommended publications
  • A Neoceratopsian Dinosaur from the Early Cretaceous of Mongolia And
    ARTICLE https://doi.org/10.1038/s42003-020-01222-7 OPEN A neoceratopsian dinosaur from the early Cretaceous of Mongolia and the early evolution of ceratopsia ✉ Congyu Yu 1 , Albert Prieto-Marquez2, Tsogtbaatar Chinzorig 3,4, Zorigt Badamkhatan4,5 & Mark Norell1 1234567890():,; Ceratopsia is a diverse dinosaur clade from the Middle Jurassic to Late Cretaceous with early diversification in East Asia. However, the phylogeny of basal ceratopsians remains unclear. Here we report a new basal neoceratopsian dinosaur Beg tse based on a partial skull from Baruunbayan, Ömnögovi aimag, Mongolia. Beg is diagnosed by a unique combination of primitive and derived characters including a primitively deep premaxilla with four pre- maxillary teeth, a trapezoidal antorbital fossa with a poorly delineated anterior margin, very short dentary with an expanded and shallow groove on lateral surface, the derived presence of a robust jugal having a foramen on its anteromedial surface, and five equally spaced tubercles on the lateral ridge of the surangular. This is to our knowledge the earliest known occurrence of basal neoceratopsian in Mongolia, where this group was previously only known from Late Cretaceous strata. Phylogenetic analysis indicates that it is sister to all other neoceratopsian dinosaurs. 1 Division of Vertebrate Paleontology, American Museum of Natural History, New York 10024, USA. 2 Institut Català de Paleontologia Miquel Crusafont, ICTA-ICP, Edifici Z, c/de les Columnes s/n Campus de la Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès Sabadell, Barcelona, Spain. 3 Department of Biological Sciences, North Carolina State University, Raleigh, NC 27695, USA. 4 Institute of Paleontology, Mongolian Academy of Sciences, ✉ Ulaanbaatar 15160, Mongolia.
    [Show full text]
  • A Dinosaur Called Tiny Free
    FREE A DINOSAUR CALLED TINY PDF Alan Durant,Jo Simpson | 32 pages | 06 Aug 2007 | HarperCollins Publishers | 9780007233908 | English | London, United Kingdom A Dinosaur Called Tiny by Alan Durant Members of its single species Compsognathus longipes could grow to around the size of a turkey. Paleontologists have found two well-preserved fossilsone in Germany in the s and the second in France more than a century later. Today, C. Many presentations still describe Compsognathus as "chicken-sized" dinosaurs because of the size of the German specimen, which is now believed to be a juvenile. Compsognathus longipes is one of the few dinosaur species whose diet is known with certainty: the remains of small, agile lizards are preserved in the bellies of both specimens. Teeth discovered in Portugal may be further fossil remains of the genus. Although not recognized as such at the time of its discovery, Compsognathus is the first theropod dinosaur known from a reasonably complete fossil skeleton. Until the s, it was the smallest-known non- avialan dinosaur, with the preceding centuries incorrectly labelling them as the closest relative of Archaeopteryx. However, dinosaurs discovered later, such as CaenagnathasiaMicroraptor and Parvicursorwere even smaller. The largest Compsognathus specimen is estimated to have weighed somewhere between A Dinosaur Called Tiny. Compsognathus were small, bipedal animals with long hind legs and longer tails, which they used for balance during locomotion. The forelimbs were smaller than the hindlimbs. The hand bore two large, clawed digits and a third, smaller A Dinosaur Called Tiny that may have been non-functional. The skull had five pairs of fenestrae skull openingsthe largest of which was for the orbit eye socket[7] with the eyes being larger in proportion to the rest of the skull.
    [Show full text]
  • Index of Subjects
    Cambridge University Press 978-1-108-47594-5 — Dinosaurs 4th Edition Index More Information INDEX OF SUBJECTS – – – A Zuul, 275 276 Barrett, Paul, 98, 335 336, 406, 446 447 Ankylosauridae Barrick, R, 383–384 acetabulum, 71, 487 characteristics of, 271–273 basal dinosauromorph, 101 acromial process, 271, 273, 487 cladogram of, 281 basal Iguanodontia, 337 actual diversity, 398 defined, 488 basal Ornithopoda, 336 adenosine diphosphate, see ADP evolution of, 279 Bates, K. T., 236, 360 adenosine triphosphate, see ATP ankylosaurids, 275–276 beak, 489 ADP (adenosine diphosphate), 390, 487 anpsids, 76 belief systems, 474, 489 advanced characters, 55, 487 antediluvian period, 422, 488 Bell, P. R., 162 aerobic metabolism, 391, 487 anterior position, 488 bennettitaleans, 403, 489 – age determination (dinosaur), 354 357 antorbital fenestra, 80, 488 benthic organisms, 464, 489 Age of Dinosaurs, 204, 404–405 Arbour, Victoria, 277 Benton, M. J., 2, 104, 144, 395, 402–403, akinetic movement, see kinetic movement Archibald, J. D., 467, 469 444–445, 477–478 Alexander, R. M., 361 Archosauria, 80, 88–90, 203, 488 Berman, D. S, 236–237 allometry, 351, 487 Archosauromorpha, 79–81, 488 Beurien, Karl, 435 altricial offspring, 230, 487 archosauromorphs, 401 bidirectional respiration, 350 Alvarez, Luis, 455 archosaurs, 203, 401 Big Al, 142 – Alvarez, Walter, 442, 454 455, 481 artifacts, 395 biogeography, 313, 489 Alvarezsauridae, 487 Asaro, Frank, 455 biomass, 415, 489 – alvarezsaurs, 168 169 ascending process of the astragalus, 488 biosphere, 2 alveolus/alveoli,
    [Show full text]
  • A Battering Ram?
    A Battering Ram? All evidence suggests that Pachycephalosaur skulls were built to withstand extreme forces 9 inches of solid bone Bone organized in a radial arrangement- structural support Articulation btw back of skull and vertebrae oriented to transfer forces linearly Articulation btw back of skull and vertebral column built to withstand sideways forces Vertebral column has tongue and groove articulations Spinal column is an S-shaped shock absorber BUT There is no ‘locking’ mechanism on skull to keep battering heads aligned Some Pachycephalosaurs have imprinted blood vessels on dome These factors suggests that head- butting may not be likely Intraspecies Competition (typically male-male) Females are typically choosey Why? Because they have more to loose Common rule in biology: Females are expensive to lose, males are cheap (e.g. deer hunting) Females choose the male most likely to provide the most successful offspring Males compete with each other for access to female vs. female chooses the strongest male Choosey females // Strong males have more offspring => SEXUAL selection Many ways to do this... But: In general, maximize competition and minimize accidental deaths (= no fitness) http://www.youtube.com/watch?v=PontCXFgs0M http://www.metacafe.com/watch/1941236/giraffe_fight/ http://www.youtube.com/watch? http://www.youtube.com/watch?v=DYDx1y38vGw http://www.youtube.com/watch?v=ULRtdk-3Yh4 Air-filled horn cores vs. solid bone skull caps... Gotta have a cheezy animated slide. Homalocephale Pachycephalosaurus Prenocephale Tylocephale Stegoceras Head butting Pachycephalosaurs Bone structure was probably strong enough to withstand collision Convex nature would favor glancing blows Instead, dome and spines seem better suited for “flank butting” So..
    [Show full text]
  • Taxonomy, Cranial Morphology, and Relationships of Parrot-Beaked Dinosaurs (Ceratopsia: Psittacosaurus)
    2 Taxonomy, Cranial Morphology, and Relationships of Parrot-Beaked Dinosaurs (Ceratopsia: Psittacosaurus) PAUL C. SERENO in 1922, well-preserved fossils of the first parrot- (Coombs 1980, 1982). For many years, Osborn’s two brief beaked dinosaur were discovered in Early Cretaceous notes on P. mongoliensis (Osborn 1923, 1924) and a descrip- horizons in the Gobi Desert of Mongolia. Now referred to tion of P. sinensis (Young 1958) provided most of the informa- a single species, Psittacosaurus mongoliensis, these remains tion available on psittacosaur morphology. include a growth series from hatchlings to adults. In sub- Recent Work. Sereno (1987) provided an overview of psit- sequent years, 15 species have been added to the genus tacosaur morphology. Portions of this dissertation were pub- Psittacosaurus and a second genus, Hongshanosaurus, was lished, including the description of two new species (P. meiley- recently described, all from Early Cretaceous rocks in ingensis, P. xinjiangensis; Sereno and Zhao 1988; Sereno et al. Asia. Although the second genus and about one-half of 1988), the synonomy of several poorly known species (Sereno the species attributed to Psittacosaurus are potentially in- 1990a), and an overview of the morphology of the clade Psit- valid, Psittacosaurus remains the most species-rich dino- tacosauridae (Sereno 1990b). Although most of this overview saurian genus, with interspecific variation concentrated can be found in You and Dodson (2004), reference is made in the skull and dentition. This paper reviews evidence only to the original source (Sereno 1990b). differentiating the named genera and species of psit- Russian psittacosaurs, including a partial skull first reported tacosaurs, outlines major cranial changes in a growth se- by Rozhdestvensky (1955, 1960) at Shestakovo in Siberia, be- ries from hatchling to adult in Psittacosaurus came the subject of a dissertation by Xijin Zhao under his mongoliensis, and provides evidence of two species direction.
    [Show full text]
  • Body-Size Evolution in the Dinosauria
    8 Body-Size Evolution in the Dinosauria Matthew T. Carrano Introduction The evolution of body size and its influence on organismal biology have received scientific attention since the earliest decades of evolutionary study (e.g., Cope, 1887, 1896; Thompson, 1917). Both paleontologists and neontologists have attempted to determine correlations between body size and numerous aspects of life history, with the ultimate goal of docu- menting both the predictive and causal connections involved (LaBarbera, 1986, 1989). These studies have generated an appreciation for the thor- oughgoing interrelationships between body size and nearly every sig- nificant facet of organismal biology, including metabolism (Lindstedt & Calder, 1981; Schmidt-Nielsen, 1984; McNab, 1989), population ecology (Damuth, 1981; Juanes, 1986; Gittleman & Purvis, 1998), locomotion (Mc- Mahon, 1975; Biewener, 1989; Alexander, 1996), and reproduction (Alex- ander, 1996). An enduring focus of these studies has been Cope’s Rule, the notion that body size tends to increase over time within lineages (Kurtén, 1953; Stanley, 1973; Polly, 1998). Such an observation has been made regarding many different clades but has been examined specifically in only a few (MacFadden, 1986; Arnold et al., 1995; Jablonski, 1996, 1997; Trammer & Kaim, 1997, 1999; Alroy, 1998). The discordant results of such analyses have underscored two points: (1) Cope’s Rule does not apply universally to all groups; and (2) even when present, size increases in different clades may reflect very different underlying processes. Thus, the question, “does Cope’s Rule exist?” is better parsed into two questions: “to which groups does Cope’s Rule apply?” and “what process is responsible for it in each?” Several recent works (McShea, 1994, 2000; Jablonski, 1997; Alroy, 1998, 2000a, 2000b) have begun to address these more specific questions, attempting to quantify patterns of body-size evolution in a phylogenetic (rather than strictly temporal) context, as well as developing methods for interpreting the resultant patterns.
    [Show full text]
  • The Dinosaur Field Guide Supplement
    The Dinosaur Field Guide Supplement September 2010 – December 2014 By, Zachary Perry (ZoPteryx) Page 1 Disclaimer: This supplement is intended to be a companion for Gregory S. Paul’s impressive work The Princeton Field Guide to Dinosaurs, and as such, exhibits some similarities in format, text, and taxonomy. This was done solely for reasons of aesthetics and consistency between his book and this supplement. The text and art are not necessarily reflections of the ideals and/or theories of Gregory S. Paul. The author of this supplement was limited to using information that was freely available from public sources, and so more information may be known about a given species then is written or illustrated here. Should this information become freely available, it will be included in future supplements. For genera that have been split from preexisting genera, or when new information about a genus has been discovered, only minimal text is included along with the page number of the corresponding entry in The Princeton Field Guide to Dinosaurs. Genera described solely from inadequate remains (teeth, claws, bone fragments, etc.) are not included, unless the remains are highly distinct and cannot clearly be placed into any other known genera; this includes some genera that were not included in Gregory S. Paul’s work, despite being discovered prior to its publication. All artists are given full credit for their work in the form of their last name, or lacking this, their username, below their work. Modifications have been made to some skeletal restorations for aesthetic reasons, but none affecting the skeleton itself.
    [Show full text]
  • Cope's Rule and the Universal Scaling Law Of
    vol. 186, no. 2 the american naturalist august 2015 Cope’s Rule and the Universal Scaling Law of Ornament Complexity Pasquale Raia,1,* Federico Passaro,1 Francesco Carotenuto,1 Leonardo Maiorino,2 Paolo Piras,2 Luciano Teresi,3 Shai Meiri,4 Yuval Itescu,4 Maria Novosolov,4 Mattia Antonio Baiano,5 Ricard Martínez,6 and Mikael Fortelius7 1. Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse (DISTAR), Università degli Studi di Napoli Federico II, Largo San Marcellino 10, 80138 Naples, Italy; 2. Dipartimento di Scienze, Università degli Studi Roma Tre, Largo San Leonardo Murialdo 1, 00146 Rome, Italy; 3. Dipartimento di Matematica e Fisica, Università degli Studi Roma Tre, Largo San Leonardo Murialdo 1, 00146 Rome, Italy; 4. Department of Zoology, Tel Aviv University, 26 Michal Street, 63261 Tel Aviv, Israel; 5. Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer Escola Industrial 23, Sabadell, E-08201 Catalonia, Spain; 6. Departament de Geologia (Paleontologia), Universitat Autónoma de Barcelona, 08193 Bellaterra, Spain; 7. Department of Geosciences and Geography, Gustaf Hällströmin katu 2a, University of Helsinki, FI-00014 Helsinki, Finland Submitted August 8, 2014; Accepted February 27, 2015; Electronically published May 29, 2015 Online enhancement: appendix. Dryad data: http://dx.doi.org/10.5061/dryad.50dr8. abstract: Luxuriant, bushy antlers, bizarre crests, and huge, twist- ular, sometimes bizarre, diversity of ornament shapes and ing horns and tusks are conventionally understood as products of sex- sizes. The evolution of these traits has always attracted ual selection. This view stems from both direct observation and from considerable interest from evolutionary biologists (Darwin the empirical finding that the size of these structures grows faster 1871).
    [Show full text]
  • 2.2.2. Ceratopsia
    Evolution der Dinosaurier Ornithischia (Vogelbecken) Ein Beitrag von Ralf Krüger 2. Cerapoda (2) 2.2.2. Ceratopsia Die Vertreter der Ceratopsia gehören zu pflanzenfressenden (herbivore) Dinosaurier. Sie besaßen häufig einen Nackenschild und /oder Hörner. Die Vorderbeine waren stets kürzer als die Hinterbeine. Die Ceratopsia enthielten die Familien der Neoceratopsia, die der Psittacosauridae und die der Ceratopsidae. 2.2.2.1. Neoceratopsia Die Neoceratopsia liefen sowohl biped als auch quadruped. Sie waren mit wenigen Ausnahmen nur 0,5 bis 3 Meter lang. Yinlong (urtümlicher Ceratopsia) Zeit: Jura 163-157 Fundort: Yinlong, China Länge: 1,2 m Höhe: 0,4 Gewicht: 30 kg Rekonstruktionsbild vom Yinglong Quelle: Nobu Tamura (CC BY-SA 3.0) Chaoyangsaurus Zeit: Jura 160-145 Fundort: Chaoyang, China Länge: 1,5 m Höhe: 0,5 m Gewicht: 40 kg Rekonstruktionsbild vom Chaoyangsaurus Quelle: Nobu Tamura (CC BY-SA 3.0) Archaeoceratops Zeit: Kreide 126-113 Fundort: Gansu, China Länge: 1,3 m Höhe: 0,4 m Gewicht: 35 kg Rekonstruktionsbild vom Archaeoceratops Quelle: Nobu Tamura (CC BY-SA 3.0) Auroraceratops Zeit: Kreide 126-113 Fundort: Gansu, China Länge: 5,5 m Höhe: 2 m Gewicht: 650 kg Rekonstruktionsbild vom Auroraceratops Quelle: Nobu Tamura (CC BY-SA 3.0) GSE-Team 2016 Seite 1 von 9 © Ralf Krüger Evolution der Dinosaurier Ornithischia (Vogelbecken) Ein Beitrag von Ralf Krüger Liaoceratops Zeit: Kreide 139-128 Fundort: Liaoning, China Länge: 1 m Höhe: 0,3 m Gewicht: 20 kg Rekonstruktionsbild vom Liaoceratops Quelle: Nobu Tamura (CC BY-SA 3.0) Yamaceratops
    [Show full text]
  • A New Basal Ornithopod Dinosaur from the Lower Cretaceous of China
    A new basal ornithopod dinosaur from the Lower Cretaceous of China Yuqing Yang1,2,3, Wenhao Wu4,5, Paul-Emile Dieudonné6 and Pascal Godefroit7 1 College of Resources and Civil Engineering, Northeastern University, Shenyang, Liaoning, China 2 College of Paleontology, Shenyang Normal University, Shenyang, Liaoning, China 3 Key Laboratory for Evolution of Past Life and Change of Environment, Province of Liaoning, Shenyang Normal University, Shenyang, Liaoning, China 4 Key Laboratory for Evolution of Past Life and Environment in Northeast Asia, Ministry of Education, Jilin University, Changchun, Jilin, China 5 Research Center of Paleontology and Stratigraphy, Jilin University, Changchun, Jilin, China 6 Instituto de Investigación en Paleobiología y Geología, CONICET, Universidad Nacional de Río Negro, Rio Negro, Argentina 7 Directorate ‘Earth and History of Life’, Royal Belgian Institute of Natural Sciences, Brussels, Belgium ABSTRACT A new basal ornithopod dinosaur, based on two nearly complete articulated skeletons, is reported from the Lujiatun Beds (Yixian Fm, Lower Cretaceous) of western Liaoning Province (China). Some of the diagnostic features of Changmiania liaoningensis nov. gen., nov. sp. are tentatively interpreted as adaptations to a fossorial behavior, including: fused premaxillae; nasal laterally expanded, overhanging the maxilla; shortened neck formed by only six cervical vertebrae; neural spines of the sacral vertebrae completely fused together, forming a craniocaudally-elongated continuous bar; fused scapulocoracoid with prominent
    [Show full text]
  • A New Taxon of Basal Ceratopsian from China and the Early Evolution of Ceratopsia
    RESEARCH ARTICLE A New Taxon of Basal Ceratopsian from China and the Early Evolution of Ceratopsia Fenglu Han1,2*, Catherine A. Forster3, James M. Clark3, Xing Xu2 1 School of Earth Sciences, China University of Geosciences, Wuhan, China, 2 Key Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China, 3 Department of Biological Sciences, The George Washington University, Washington, District of Columbia, United States of America * [email protected] a11111 Abstract Ceratopsia is one of the best studied herbivorous ornithischian clades, but the early evolu- tion of Ceratopsia, including the placement of Psittacosaurus, is still controversial and unclear. Here, we report a second basal ceratopsian, Hualianceratops wucaiwanensis gen. OPEN ACCESS et sp. nov., from the Upper Jurassic (Oxfordian) Shishugou Formation of the Junggar Basin, Citation: Han F, Forster CA, Clark JM, Xu X (2015) A northwestern China. This new taxon is characterized by a prominent caudodorsal process New Taxon of Basal Ceratopsian from China and the Early Evolution of Ceratopsia. PLoS ONE 10(12): on the subtemporal ramus of the jugal, a robust quadrate with an expansive quadratojugal e0143369. doi:10.1371/journal.pone.0143369 facet, a prominent notch near the ventral region of the quadrate, a deep and short dentary, Editor: Matthew Shawkey, University of Akron, and strongly rugose texturing on the lateral surface of the dentary. Hualianceratops shares UNITED STATES several derived characters with both Psittacosaurus and the basal ceratopsians Yinlong, Received: July 15, 2015 Chaoyangsaurus, and Xuanhuaceratops. A new comprehensive phylogeny of ceratopsians weakly supports both Yinlong and Hualianceratops as chaoyangsaurids (along with Accepted: November 3, 2015 Chaoyangsaurus and Xuanhuaceratops), as well as the monophyly of Chaoyangosauridae Published: December 9, 2015 + Psittacosaurus.
    [Show full text]
  • 1 PHILATELIC SUPPLIES (M.B.O'neill)
    List up-dated Winter 2020 (10.12.20) P R E H I S T O R Y (1 9 6 5 - 2 0 2 0) SPECIAL OFFERS EDITION (includes Fossils and Prehistoric Animals) PHILATELIC SUPPLIES (M.B.O'Neill) 359 Norton Way South Letchworth Garden City HERTS ENGLAND SG6 1SZ Telephone 01462-684191 during office hours 9.15-3.15pm Mon.-Fri.) Web-site: www.philatelicsupplies.co.uk email: [email protected] TERMS OF BUSINESS: & Notes on these lists: (Please read before ordering). 1). All stamps are unmounted mint unless specified otherwise. Our prices are also in Sterling, (£). 2). Lists are updated about every 26 weeks to include most recent stock movements and New Issues; they are therefore reasonably accurate stockwise 100% pricewise. This reduces the need for "credit notes" and refunds. Alternatives may be listed in case some items are out of stock. However, these popular lists are still best used as soon as possible. Next Prehistory listings will be printed in 6 & 12 months time so please indicate when next we should send a list on your order form. 3). New Issues Services can be provided if you wish to keep your collection up to date on a Standing Order basis. Details of this service & forms can be sent on request. Regret we do not run an on approval service. 4).All orders on our order forms are attended to by return of post. We will keep a photocopy it and return your annotated original. 5). Other Thematic Lists are available on request; Minerals/Geology, Fauna, 6).
    [Show full text]