The Hypothesis of Saurischian Pneumaticity

Total Page:16

File Type:pdf, Size:1020Kb

The Hypothesis of Saurischian Pneumaticity Geological Society, London, Special Publications Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurischian pneumaticity Darren Naish Geological Society, London, Special Publications 2010; v. 343; p. 229-236 doi:10.1144/SP343.13 Email alerting click here to receive free email alerts when new articles cite this service article Permission click here to seek permission to re-use all or part of this article request Subscribe click here to subscribe to Geological Society, London, Special Publications or the Lyell Collection Notes Downloaded by University of Portsmouth on 6 October 2010 © 2010 Geological Society of London Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurischian pneumaticity DARREN NAISH School of Earth & Environmental Sciences, Burnaby Building, Burnaby Road, University of Portsmouth, Portsmouth PO1 3QL, UK (e-mail: [email protected]) Abstract: Saurischian dinosaurs were pneumatic animals. The presence of invasive skeletal for- amina leading to large internal chambers within the skeleton strongly indicate the presence of avian-style skeletal pneumaticity of the skeleton in sauropodomorphs and non-avian theropods. While the hypothesis of skeletal pneumaticity has undergone a renaissance in recent years, it was initially promoted during the late 1800s after dinosaur fossils from the English Lower Cretac- eous Wealden Supergroup led Richard Owen and Harry Seeley to note the pneumatic, bird-like fea- tures of the vertebrae they described (Hermann von Meyer had also briefly alluded to skeletal pneumaticity in dinosaurs during the 1830s). In describing the theropod Becklespinax altispinax from the Hastings Beds Group (at the time referred to Megalosaurus), Richard Owen proposed that the laminae on the neural arch served to house ‘parts of the lungs’. He evidently imagined Becklespinax to exhibit avian-style post-cranial skeletal pneumaticity. In 1870 Harry Seeley described two sauropod vertebrae from the Wealden Supergroup, naming them Ornithopsis hulkei. Contrary to what is often stated, Seeley did not identify Ornithopsis as a pterosaur, but as an animal that might ‘bridge over’ the gap between birds and pterosaurs, while at the same time having some affinity with dinosaurs. The lateral foramina and internal bony cavities of one of these specimens were regarded by Seeley as allowing ‘the prolongation of the peculiarly avian respiratory system into the bones’, and he emphasized ‘the lightest and airiest plan’ of the specimen. In 1876 Owen described the Wessex Formation sauropod Chondrosteosaurus gigas. While regarding the lateral fossae as probably having ‘lodged a saccular process of the lung’, Owen now took the opportunity to attack Seeley’s claims of pneumaticity in Ornithopsis, arguing that the internal cavities in Chondrosteosaurus ‘were occupied in the living reptile by unossified cartilage, or chondrine’. The name Chondrosteosaurus gigas (‘giant cartilage and bone lizard’) also looks like a direct assault on Seeley’s proposal of a pneumatic vertebral interior. Owen’s actions seem odd given that he was familiar with the internal morphology of avian ver- tebrae (which are often strikingly similar to those of sauropods). However, both authors have proved insighful in correctly identifying skeletal pneumaticity during this early phase of dinosaur research. A thorough historical review of early ideas on dinosaurian pneumaticity is still required. In terms of its significance for early dinosaur discov- unnamed) tetanuran theropod of uncertain affinities eries, the Lower Cretaceous Wealden Supergroup of (Benson et al. 2009). southern England must rank as one of the most The term ‘Wealden’ refers to a series of non- important geological units. It yielded Mantell’s marine mudstones, sandstones and other strata that original Iguanodon material during the 1820s, the were deposited in two sub-basins located in what armoured dinosaur Hylaeosaurus armatus during is now SE England: the Weald sub-basin of the the 1830s, the earliest sauropod discoveries during English mainland; and the Wessex sub-basin of the 1840s, and what proved to be a pivotal form in the Isle of Wight and Dorset (Martill & Naish early ideas on the evolutionary relationship 2001; Radley 2004, 2006a, b). While the strata of between dinosaurs and birds, Hypsilophodon,in both the Weald and Wessex sub-basins were pre- 1869. Despite the fact that Wealden exposures viously referred to as ‘the Wealden Group’, they have been well explored and extensively studied are now known as the Wealden Supergroup since the early 1800s, they continue to yield new (Fig. 1). Within the Weald sub-basin, the oldest dinosaurs, with recently described taxa including unit is the Berriasian–Valanginian Hastings Beds the spinosauroid Baryonyx walkeri (Charig & Group. Younger than the Hastings Beds Group, Milner 1986), the ankylosaur Polacanthus rudg- but also occurring within the Weald sub-basin, is wickensis (Blows 1996), the allosauroid Neovenator the Weald Clay Group: this unit is mostly Hauteri- salerii (Hutt et al. 1996), the basal tyrannosauroid vian and Barremian, but might extend into the Eotyrannus lengi (Hutt et al. 2001), the extremely Aptian as well (Allen & Wimbledon 1991). unusual neosauropod Xenoposeidon proneneukos Finally, within the Wessex sub-basin, the Wealden (Taylor & Naish 2007) and a large (as yet Group (sensu stricto) is mostly Barremian and From:Moody, R. T. J., Buffetaut, E., Naish,D.&Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications, 343, 229–236. DOI: 10.1144/SP343.13 0305-8719/10/$15.00 # The Geological Society of London 2010. 230 D. NAISH phylogeny and function of pneumaticity, there are indications that its presence may correlate not only with pulmonary structure and function but also with metabolism and growth rates (Bonde & Chris- tiansen 2003; Wedel 2003b; O’Connor 2006). A thorough review on the history of thoughts about saurischian pneumaticity has yet to appear. Here, I examine the role that Wealden Supergroup dinosaurs had in early ideas on skeletal pneumaticity. Institutional abbreviations: HASMG, Hastings Museum and Art Gallery, Hastings, UK; NHM UK, Natural History Museum, London, UK (formerly the British Musuem (Natural History)). Fig. 1. Stratigraphic diagram showing the approximate correlations between the Weald sub-basin units of the English mainland and the Wessex sub-basin units of the Isle of Wight and Dorset. Based on Kerth and Hailwood Becklespinax, the ‘first’ pneumatic (1988), Allen and Wimbledon (1991), Martill & Naish theropod (2001) and Radley (2004, 2006a, b). Among the few theropods that are known from the Hastings Beds Group is NHMUK R1828, currently extends into the Aptian (Kerth & Hailwood 1988). It known as Becklespinax altispinax (Paul 1988). Rep- includes the Wessex and Vectis formations, both of resented only by three articulated dorsal vertebrae which crop out on the Isle of Wight. discovered by Samuel Beckles some time prior to While the Wealden Supergroup is often noted as 1855 (Owen 1855), these three vertebrae are all an important unit for discoveries that have shed new that we have of this dinosaur. However, in describ- light on dinosaur diversity, less well appreciated is ing the specimen, Owen (1856) referred to some that dinosaurs from the Wealden have also proved other material (another two vertebrae and two important in terms of shaping our views on dinosaur ribs) that apparently belonged to it but have since palaeobiology. Among the most interesting and been lost. The proximal end of the robust right vexing, and arguably most important, aspect of tibia HASM G.378 from the Hastings Beds Group, dinosaur palaeobiology is the fact that saurischians identified as Allosauroidea indet., was suggested (and not ornithischians so far as we know) exhibited by Naish (2003) to perhaps be referable to B. altis- skeletal pneumaticity: a system of air sacs and pneu- pinax. It remains impossible to evaluate this in the matic diverticula were present in at least some of the absence of better B. altispinax remains. vertebrae, with basal forms exhibiting shallow Like many Wealden Supergroup dinosaurs, B. pneumatic fossae on their vertebral centra and altispinax has had a tumultuous nomenclatural derived forms possessing internalized pneumatic history, and I have largely avoided this area here cavities connected to foramina located within the (a full revision of B. altispinax is in preparation). fossae (Britt 1993; Wedel 2003a, b, 2004, 2007; However, Rauhut (2000) argued that Huene (1923) O’Connor 2006). Pterosaurs also exhibited skeletal proposed the generic name Altispinax for NHMUK pneumaticity, raising the possibility that it was R1828, and not for the indeterminate tooth regarded ancestral for ornithodirans and secondarily lost in as the type of Megalosaurus dunkeri by Kuhn ornithischians (Bonde & Christiansen 2003; Butler (1939) and Olshevsky (1991). Rauhut therefore con- et al. 2009). Skeletal pneumaticity has also been cluded that B. altispinax should be referred to as inferred for some basal archosauriforms (Erythrosu- Altispinax altispinax. This can be interpreted as chus africanus) and crocodile-group archosaurs, contradicting key statements in the literature. On which could suggest that it was primitive for naming Altispinax, Huene (1923) included Megalo- crown-group archosaurs or even for a more inclus- saurus dunkeri and M. oweni within the genus. He ive clade
Recommended publications
  • Dino Hunt Checklist Card Name Type Rarity Acanthopholis
    Dino Hunt Checklist Card Name Type Rarity Acanthopholis Dinosaur Common Acrocanthosaurus Dinosaur Rare Albertosaurus Dinosaur Rare Albertosaurus Dinosaur Ultra Rare Alioramus Dinosaur Rare Allosaurus Dinosaur Rare* Altispinax Dinosaur Rare* Amargasaurus Dinosaur Uncommon* Ammosaurus Dinosaur Uncommon* Anatotitan Dinosaur Common Anchiceratops Dinosaur Common Anchisaurus Dinosaur Common* Ankylosaurus Dinosaur Uncommon* Antarctosaurus Dinosaur Common Apatosaurus Dinosaur Uncommon* Archaeopteryx Dinosaur Rare* Archelon Dinosaur Rare Arrhinoceratops Dinosaur Common Avimimus Dinosaur Common Baby Ankylosaur Dinosaur Common Baby Ceratopsian Dinosaur Common Baby Hadrosaur Dinosaur Common Baby Raptor Dinosaur Rare Baby Sauropod Dinosaur Common Baby Theropod Dinosaur Rare Barosaurus Dinosaur Uncommon Baryonyx Dinosaur Rare* Bellusaurus Dinosaur Common Brachiosaurus Dinosaur Rare* Brachyceratops Dinosaur Uncommon Camarasaurus Dinosaur Common Camarasaurus Dinosaur Ultra Rare Camptosaurus Dinosaur Common Carnotaurus Dinosaur Rare Centrosaurus Dinosaur Common Ceratosaurus Dinosaur Rare* Cetiosaurus Dinosaur Common* Changdusaurus Dinosaur Common Chasmosaurus Dinosaur Common Chilantaisaurus Dinosaur Rare Coelophysis Dinosaur Uncommon* Coloradisaurus Dinosaur Common* Compsognathus Dinosaur Rare* Corythosaurus Dinosaur Common* Cryolophosaurus Dinosaur Rare Cynognathus Dinosaur Rare Dacentrurus Dinosaur Common* Daspletosaurus Dinosaur Rare Datousaurus Dinosaur Common Deinocheirus Dinosaur Rare* Deinonychus Dinosaur Uncommon* Deinosuchus Dinosaur Rare* Diceratops
    [Show full text]
  • Brains and Intelligence
    BRAINS AND INTELLIGENCE The EQ or Encephalization Quotient is a simple way of measuring an animal's intelligence. EQ is the ratio of the brain weight of the animal to the brain weight of a "typical" animal of the same body weight. Assuming that smarter animals have larger brains to body ratios than less intelligent ones, this helps determine the relative intelligence of extinct animals. In general, warm-blooded animals (like mammals) have a higher EQ than cold-blooded ones (like reptiles and fish). Birds and mammals have brains that are about 10 times bigger than those of bony fish, amphibians, and reptiles of the same body size. The Least Intelligent Dinosaurs: The primitive dinosaurs belonging to the group sauropodomorpha (which included Massospondylus, Riojasaurus, and others) were among the least intelligent of the dinosaurs, with an EQ of about 0.05 (Hopson, 1980). Smartest Dinosaurs: The Troodontids (like Troödon) were probably the smartest dinosaurs, followed by the dromaeosaurid dinosaurs (the "raptors," which included Dromeosaurus, Velociraptor, Deinonychus, and others) had the highest EQ among the dinosaurs, about 5.8 (Hopson, 1980). The Encephalization Quotient was developed by the psychologist Harry J. Jerison in the 1970's. J. A. Hopson (a paleontologist from the University of Chicago) did further development of the EQ concept using brain casts of many dinosaurs. Hopson found that theropods (especially Troodontids) had higher EQ's than plant-eating dinosaurs. The lowest EQ's belonged to sauropods, ankylosaurs, and stegosaurids. A SECOND BRAIN? It used to be thought that the large sauropods (like Brachiosaurus and Apatosaurus) and the ornithischian Stegosaurus had a second brain.
    [Show full text]
  • A Revised Taxonomy of the Iguanodont Dinosaur Genera and Species
    ARTICLE IN PRESS + MODEL Cretaceous Research xx (2007) 1e25 www.elsevier.com/locate/CretRes A revised taxonomy of the iguanodont dinosaur genera and species Gregory S. Paul 3109 North Calvert Station, Side Apartment, Baltimore, MD 21218-3807, USA Received 20 April 2006; accepted in revised form 27 April 2007 Abstract Criteria for designating dinosaur genera are inconsistent; some very similar species are highly split at the generic level, other anatomically disparate species are united at the same rank. Since the mid-1800s the classic genus Iguanodon has become a taxonomic grab-bag containing species spanning most of the Early Cretaceous of the northern hemisphere. Recently the genus was radically redesignated when the type was shifted from nondiagnostic English Valanginian teeth to a complete skull and skeleton of the heavily built, semi-quadrupedal I. bernissartensis from much younger Belgian sediments, even though the latter is very different in form from the gracile skeletal remains described by Mantell. Currently, iguanodont remains from Europe are usually assigned to either robust I. bernissartensis or gracile I. atherfieldensis, regardless of lo- cation or stage. A stratigraphic analysis is combined with a character census that shows the European iguanodonts are markedly more morpho- logically divergent than other dinosaur genera, and some appear phylogenetically more derived than others. Two new genera and a new species have been or are named for the gracile iguanodonts of the Wealden Supergroup; strongly bipedal Mantellisaurus atherfieldensis Paul (2006. Turning the old into the new: a separate genus for the gracile iguanodont from the Wealden of England. In: Carpenter, K. (Ed.), Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs.
    [Show full text]
  • Chapter 10, the Mistaken Extinction, by Lowell Dingus and Timothy Rowe, New York, W
    Chapter 10, The Mistaken Extinction, by Lowell Dingus and Timothy Rowe, New York, W. H. Freeman, 1998. CHAPTER 10 Dinosaurs Challenge Evolution Enter Sir Richard Owen More than 150 years ago, the great British naturalist Richard Owen (fig. 10.01) ignited the controversy that Deinonychus would eventually inflame. The word "dinosaur" was first uttered by Owen in a lecture delivered at Plymouth, England in July of 1841. He had coined the name in a report on giant fossil reptiles that were discovered in England earlier in the century. The root, Deinos, is usually translated as "terrible" but in his report, published in 1842, Owen chose the words "fearfully great"1. To Owen, dinosaurs were the fearfully great saurian reptiles, known only from fossil skeletons of huge extinct animals, unlike anything alive today. Fig. 10.01 Richard Owen as, A) a young man at about the time he named Dinosauria, B) in middle age, near the time he described Archaeopteryx, and C) in old age. Dinosaur bones were discovered long before Owen first spoke their name, but no one understood what they represented. The first scientific report on a dinosaur bone belonging was printed in 1677 by Rev. Robert Plot in his work, The Natural History of Oxfordshire. This broken end of a thigh bone, came to Plot's attention during his research. It was nearly 60 cm in circumference--greater than the same bone in an elephant (fig.10.02). We now suspect that it belonged to Megalosaurus bucklandii, a carnivorous dinosaur now known from Oxfordshire. But Plot concluded that it "must have been a real Bone, now petrified" and that it resembled "exactly the figure of the 1 Chapter 10, The Mistaken Extinction, by Lowell Dingus and Timothy Rowe, New York, W.
    [Show full text]
  • New Perspectives on Pterosaur Palaeobiology
    Downloaded from http://sp.lyellcollection.org/ by guest on September 24, 2021 New perspectives on pterosaur palaeobiology DAVID W. E. HONE1*, MARK P. WITTON2 & DAVID M. MARTILL2 1Queen Mary University of London, Mile End Road, London E14NS, UK 2School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Road, Portsmouth PO1 3QL, UK *Correspondence: [email protected] Abstract: Pterosaurs were the first vertebrates to evolve powered flight and occupied the skies of the Mesozoic for 160 million years. They occurred on every continent, evolved their incredible pro- portions and anatomy into well over 100 species, and included the largest flying animals of all time among their ranks. Pterosaurs are undergoing a long-running scientific renaissance that has seen elevated interest from a new generation of palaeontologists, contributions from scientists working all over the world and major advances in our understanding of their palaeobiology. They have espe- cially benefited from the application of new investigative techniques applied to historical speci- mens and the discovery of new material, including detailed insights into their fragile skeletons and their soft tissue anatomy. Many aspects of pterosaur science remain controversial, mainly due to the investigative challenges presented by their fragmentary, fragile fossils and notoriously patchy fossil record. With perseverance, these controversies are being resolved and our understand- ing of flying reptiles is increasing. This volume brings together a diverse set of papers on numerous aspects of the biology of these fascinating reptiles, including discussions of pterosaur ecology, flight, ontogeny, bony and soft tissue anatomy, distribution and evolution, as well as revisions of their taxonomy and relationships.
    [Show full text]
  • 120. Wealden Greensand Area Profile: Supporting Documents
    National Character 120. Wealden Greensand Area profile: Supporting documents www.naturalengland.org.uk 1 National Character 120. Wealden Greensand Area profile: Supporting documents Introduction National Character Areas map As part of Natural England’s responsibilities as set out in the Natural Environment 1 2 3 White Paper , Biodiversity 2020 and the European Landscape Convention , we are North revising profiles for England’s 159 National Character Areas (NCAs). These are areas East that share similar landscape characteristics, and which follow natural lines in the landscape rather than administrative boundaries, making them a good decision- Yorkshire making framework for the natural environment. & The North Humber NCA profiles are guidance documents which can help communities to inform their West decision-making about the places that they live in and care for. The information they contain will support the planning of conservation initiatives at a landscape East scale, inform the delivery of Nature Improvement Areas and encourage broader Midlands partnership working through Local Nature Partnerships. The profiles will also help West Midlands to inform choices about how land is managed and can change. East of England Each profile includes a description of the natural and cultural features that shape our landscapes, how the landscape has changed over time, the current key London drivers for ongoing change, and a broad analysis of each area’s characteristics and ecosystem services. Statements of Environmental Opportunity (SEOs) are South East suggested, which draw on this integrated information. The SEOs offer guidance South West on the critical issues, which could help to achieve sustainable growth and a more secure environmental future.
    [Show full text]
  • Geotourism and Geoconservation on the Isle of Wight, UK: Balancing Science with Commerce
    Carpenter: Rocky start of Dinosaur National Monument… Geoconservation Research Original Article Geotourism and Geoconservation on the Isle of Wight, UK: Balancing Science with Commerce Martin I. Simpson Lansdowne, Military Rd, Chale, Isle of Wight, PO38 2HH, UK. Abstract The Isle of Wight has a rich and varied geological heritage which attracts scientists, tourists and fossil collectors, both private and commercial. Each party has a role to play in geoconservation and geotourism, but a policy on the long term curation of scientifically important specimens is essential to prevent future conflicts. A new code of conduct is recommended, based on the one adopted on the Jurassic Coast of Dorset. I have spent over 40 years living on the Island and working in the tourist industry running geology field-trips for both academics and tourists, and managing one of the longest running geological gift shops. I see the geological heritage and fossil sites as valuable geotourism assets, and envisage no problems with respect to the scientifically important material provided that a clear collecting policy is adopted, and the local museum generates funding to ensure that significant finds remain on the Island. A positive attitude is recommended in view of past experiences. Corresponding Author: Martin I. Simpson Lansdowne, Military Rd, Chale, Isle of Wight, PO38 2HH, UK. Email: [email protected] Keywords: Palaeontology, Geology, Isle of Wight, Tourism. Introduction with few formations absent, probably one of the best successions of this type in Europe (Fig 1c).Once a larger landmass joined to the mainland The Isle of Wight is a small, vaguely lozenge-shaped island situated as recently as 9000 years ago, what remains as 'Wight Island' or 'Vecta just off the central south coast of England, about 113 km south west of Insula' is eroding away at a rate of one metre per year in places (Munt London (Fig 1a, 1b), renowned for its balmy climate and golden, sandy 2016), but this erosion has produced spectacular scenery and iconic beaches.
    [Show full text]
  • Back Matter (PDF)
    Index Note: Page numbers in italic denote figures. Page numbers in bold denote tables. Abel, Othenio (1875–1946) Ashmolean Museum, Oxford, Robert Plot 7 arboreal theory 244 Astrodon 363, 365 Geschichte und Methode der Rekonstruktion... Atlantosaurus 365, 366 (1925) 328–329, 330 Augusta, Josef (1903–1968) 222–223, 331 Action comic 343 Aulocetus sammarinensis 80 Actualism, work of Capellini 82, 87 Azara, Don Felix de (1746–1821) 34, 40–41 Aepisaurus 363 Azhdarchidae 318, 319 Agassiz, Louis (1807–1873) 80, 81 Azhdarcho 319 Agustinia 380 Alexander, Annie Montague (1867–1950) 142–143, 143, Bakker, Robert. T. 145, 146 ‘dinosaur renaissance’ 375–376, 377 Alf, Karen (1954–2000), illustrator 139–140 Dinosaurian monophyly 93, 246 Algoasaurus 365 influence on graphic art 335, 343, 350 Allosaurus, digits 267, 271, 273 Bara Simla, dinosaur discoveries 164, 166–169 Allosaurus fragilis 85 Baryonyx walkeri Altispinax, pneumaticity 230–231 relation to Spinosaurus 175, 177–178, 178, 181, 183 Alum Shale Member, Parapsicephalus purdoni 195 work of Charig 94, 95, 102, 103 Amargasaurus 380 Beasley, Henry Charles (1836–1919) Amphicoelias 365, 366, 368, 370 Chirotherium 214–215, 219 amphisbaenians, work of Charig 95 environment 219–220 anatomy, comparative 23 Beaux, E. Cecilia (1855–1942), illustrator 138, 139, 146 Andrews, Roy Chapman (1884–1960) 69, 122 Becklespinax altispinax, pneumaticity 230–231, Andrews, Yvette 122 232, 363 Anning, Joseph (1796–1849) 14 belemnites, Oxford Clay Formation, Peterborough Anning, Mary (1799–1847) 24, 25, 113–116, 114, brick pits 53 145, 146, 147, 288 Benett, Etheldred (1776–1845) 117, 146 Dimorphodon macronyx 14, 115, 294 Bhattacharji, Durgansankar 166 Hawker’s ‘Crocodile’ 14 Birch, Lt.
    [Show full text]
  • Lyons SCIENCE 2021 the Influence of Juvenile Dinosaurs SUPPL.Pdf
    science.sciencemag.org/content/371/6532/941/suppl/DC1 Supplementary Materials for The influence of juvenile dinosaurs on community structure and diversity Katlin Schroeder*, S. Kathleen Lyons, Felisa A. Smith *Corresponding author. Email: [email protected] Published 26 February 2021, Science 371, 941 (2021) DOI: 10.1126/science.abd9220 This PDF file includes: Materials and Methods Supplementary Text Figs. S1 and S2 Tables S1 to S7 References Other Supplementary Material for this manuscript includes the following: (available at science.sciencemag.org/content/371/6532/941/suppl/DC1) MDAR Reproducibility Checklist (PDF) Materials and Methods Data Dinosaur assemblages were identified by downloading all vertebrate occurrences known to species or genus level between 200Ma and 65MA from the Paleobiology Database (PaleoDB 30 https://paleobiodb.org/#/ download 6 August, 2018). Using associated depositional environment and taxonomic information, the vertebrate database was limited to only terrestrial organisms, excluding amphibians, pseudosuchians, champsosaurs and ichnotaxa. Taxa present in formations were confirmed against the most recent available literature, as of November, 2020. Synonymous taxa or otherwise duplicated taxa were removed. Taxa that could not be identified to genus level 35 were included as “Taxon X”. GPS locality data for all formations between 200MA and 65MA was downloaded from PaleoDB to create a minimally convex polygon for each possible formation. Any attempt to recreate local assemblages must include all potentially interacting species, while excluding those that would have been separated by either space or time. We argue it is 40 acceptable to substitute formation for home range in the case of non-avian dinosaurs, as range increases with body size.
    [Show full text]
  • Phylogeny and Avian Evolution Phylogeny and Evolution of the Aves
    Phylogeny and Avian Evolution Phylogeny and Evolution of the Aves I. Background Scientists have speculated about evolution of birds ever since Darwin. Difficult to find relatives using only modern animals After publi cati on of “O rigi i in of S peci es” (~1860) some used birds as a counter-argument since th ere were no k nown t ransiti onal f orms at the time! • turtles have modified necks and toothless beaks • bats fly and are warm blooded With fossil discovery other potential relationships! • Birds as distinct order of reptiles Many non-reptilian characteristics (e.g. endothermy, feathers) but really reptilian in structure! If birds only known from fossil record then simply be a distinct order of reptiles. II. Reptile Evolutionary History A. “Stem reptiles” - Cotylosauria Must begin in the late Paleozoic ClCotylosauri a – “il”“stem reptiles” Radiation of reptiles from Cotylosauria can be organized on the basis of temporal fenestrae (openings in back of skull for muscle attachment). Subsequent reptilian lineages developed more powerful jaws. B. Anapsid Cotylosauria and Chelonia have anapsid pattern C. Syypnapsid – single fenestra Includes order Therapsida which gave rise to mammalia D. Diapsida – both supppratemporal and infratemporal fenestrae PttPattern foun did in exti titnct arch osaurs, survi iiving archosaurs and also in primitive lepidosaur – ShSpheno don. All remaining living reptiles and the lineage leading to Aves are classified as Diapsida Handout Mammalia Extinct Groups Cynodontia Therapsida Pelycosaurs Lepidosauromorpha Ichthyosauria Protorothyrididae Synapsida Anapsida Archosauromorpha Euryapsida Mesosaurs Amphibia Sauria Diapsida Eureptilia Sauropsida Amniota Tetrapoda III. Relationshippp to Reptiles Most groups present during Mesozoic considere d ancestors to bird s.
    [Show full text]
  • Evolution of the Caudal Vertebral Series in Macronarian Sauropod Dinosaurs: Morphofunctional and Phylogenetic Implications
    Evolution of the Caudal Vertebral Series in Macronarian Sauropod Dinosaurs: Morphofunctional and Phylogenetic Implications A Thesis Submitted to the Faculty of Drexel University By Lucio Manuel Ibiricu In partial fulfillment of the Requirements for the degree of Doctor of Philosophy October 2010 © Copyright 2010 Lucio Manuel Ibiricu. All Rights Reserved ii DEDICATION To the people who trusted me and my grandfather (Manolo) iii ACKNOWLEDGMENTS First of all, I want to thank my advisor, Dr. Lacovara, for several things: for admitting me in the program, for supporting me constantly, especially at the beginning of my experience in the United States and for exposing me to ―big‖ people in Paleontology. Ken gave me the opportunity to work and learn; I will always be grateful to him. I also thank Dr. Spotila, the head of my committee, for always supporting me and for helping in the organization, always very important during the PhD. I really appreciate it. In addition, I want to thank Dr. Dodson for accepting to be part of my committee. His presence is a privilege to me and it is an honor to say that he was part of my PhD committee. I would like to express my gratitude to Dr. Gallagher. He taught me the majority of my first courses at Drexel and I always learned a lot from him. Also, my first TA was under his supervision; therefore, I want to thank him for all the support he gave me both as student and as a TA. Likewise, I would like to thank Dr. Kilham, because her presence in my committee pushed me to learn more about ecology, which helped me integrate dinosaurs and their environment.
    [Show full text]
  • THE GENERA of REPTILES. By
    T he Genera of Re pt il e s. By BARON FRANCIS NOPCSA (Budapest). (Eingelangt am 11. Mai 1927.) Among all the Paleontologists living none has dealt with the recent and fossil reptiles with a wider grasp than Prof. L. D o l l o and at his anniversary it seems quite appropriate to review their whole array. The classification used in the following enumeration of all genera of reptiles is much the same as in my hook „Die Familien der Reptilien“; at this instance however an effort has been made to give a precise definition of every systematic unit. Alteration of the classification became necessary among the Dino- cephalians, the Nothosaurians, the Lacertilians (called here Sauroidea), the Coelurosauroidea and the Crocodilia. The new classification of the Sauroidea is intermediate between the classifications proposed by B o u l e n g e r and C a m p . Naturally as a rule only those fossil genera are referred to, that are more than simple catalogue numbers that facilitate the finding of the respective piece in a collection; indeterminable problematic genera of small interest have mostly been omitted. Extinct units and genera are marked in the lists with a cross ("j*). Difficulties of classification have been encountered in the Lacertilia and Ophidia, for in the new system the conventional families of recent Lizards and Serpents has been given subfamily rank and conse­ quently the different recent subfamilies had to be dropped. Instead of these subfamilies the minor units were separated by greek letters. — The tedious and complicated revision of the genera of recent Lizards and Snakes was done by Prof.
    [Show full text]