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THE CONODONTA G- lesources,and Morphology,Taxonomy, Paleoecology, andEvolutionary History of a Long-ExtinctAnimal Phylum 3 azI High-pressure WALTERC.SWEEL EE: TheOhio State University fIt;:t F by the -i-:ztlog' fr, :.1 a olution- b.-;:n- New York . Oxford CLARENDON PRESS. OXFORD . 1988 PREFACE Conodonts are common fossils.Almost anyone portant group, I ofer my account of the who dealson a regularbasis with Paleozoicand Conodonta. Triassic marine rocks has probably seena few For their "witting" or unwitting contribu- of them. Through the last 30 yedrs conodonts tions to what I believe I know about the fasci- have come to be of exceptionalvalue in bio- nating gtoup of extinct animals described on stratigraphy, and they now have pride of place the following pages,I am gateful to a long list as index fossils in many parts of the geologic of my students and faculty colleaguesat The column. But conodonts are extinct and are un- Ohio StateUniversity, especiallyStig M. Bergs- knowo to most neontologists. The sketchiness trijm, and to members of the Pander Society, ofinformation about them in most texts on in- an intemational group of exceptionally good- vertebrate paleontology may result either fiom natured "conodontologists"that has met fre- the fact that they are microfossils that haye quently and infomally thougl the last 20 only lately come to be important as strati- years to share infomation about conodonts, graphic tools, or from the fact that nothing argueconclusions, and correct the misappre- quite like them exists today, so their zoologic hensionsof its seniormembers. relations axeuncertain. Karen Tyler, faculty illustrator at The Ohio In this monograph, I provide a summary of State University, drafted nea y all the figures information about a group with which I have from my very crude copy and assistedwith la- worked all my adult life. Charts that name con- beling others. Dr. Jerzy Dzik, of the Polish odont-based biozones and show stratigraphic Acaderny of Sciences, Warszawa, provided rangeofselected conodont speciesare included about half the stippled drawings of conodonts as Appendix B. However, I have purposely that gace various figuresin Chapter 5- The an- avoided a recap of conodont biostratigtaphy istry of tlrese two good friends is plainly evi- becauseit is constantly changing and cunent dent in their work and is warmlY views are readily available in various other acknowledged. places.Instead, I focus here on the conodonts I am also grateful to Sue Shipley and David as a group of extinct animals, about which it is Little, of The Ohio State University, for their important to know as much as possible be- help in completing various parts of the manu- fore assessing their distribution biostrati- script and illustrations, and to Mark Klefner, graphically. who graciously cornpiled information on the Of course, one takes considerablerisk in at- ranges of Silurian conodonts and assembled tempting such a summary, particularly of a the Silurian chart in Appendix B. The Depart- group of animals known only from its fossil re- ment of Geology and Mineralology generously cord, becausemost of what I think I know assumedmuch of the expenseof drafting and about conodontsas animals is either conjecture photogaphy. or a highly personalinterpretation ofa still-irn- perfect fossil record. So, with the caveal that Columbus, Ohio w.c.s. whal followsis only one way of viewingan im- February 1988 CONTENTS l. Inhoduction l.l History ofdiscovery and study 3 1.2 Achievements 7 1.3 Pending problems 8 2. Skeletal anatomy 11 2.1 Composition of conodont elements ll 2.2 Structure of skeletalelements t2 2.3 Shapesof element crowns t4 2.3.1 Coniform crowns l5 2.3.2 Ramiform crowns t6 2.3.3 Rastratecrowns l8 2.3.4 Pectiniformcrowns t9 2.4 Symmetry- and curvature-transitionseries 22 2.5 Skeletalapparatuses 2.6 Symmetry of elements,element pairs, and apparatuses 25 J. Whole-animal anatomy 28 3.1 The Scottish Carboniferousspecimens 28 3.2 The Waukeshaspecimen 3Z 3.3 Histology of demineralizedtissues 33 3.4 Summary JJ 4. Taxonomy 35 4.1 Form taxonomy J) 4.2 Multielementtaxonomy 36 4.2.1 Multielementmethodology 4-3 Multielement classifications 4l 4.4 A revised multielement classification 41 5. The major conodontgroups 45 5.1 Introduction 45 5.2 Cavidonti and Conodonti 45 5.3 The Proconodontida (Cavidonti) and its families (Fig. 5.1) 45 5.4 The Belodellida and its families (Fig. 5.1) 49 5.5 OrderProtopanderodontida, new 5l 5.5.1 Family ProtopanderodontidaeLindstrtim, 1970 52 5.5.2 Family ClavohamulidaeLindstrdm, 1970 53 5.5.3 Family AcanthodontidaeLindstritm, 1970 54 5.5.4 Family DrepanoistodontidaeFihraeus and Nowlan, 1978 54 t vtl III eIc,(Ju?rrr^opro ele-I pue opprw z.€.9 B €€l II pue I selci(c I'9 9 B €[l selc,brsplo-puoces t.g ) r'8 ztr selc^crurel-?uo'I Z'2 9 t ztr uroll?dIpre^O lZ'9 I 0€r surened./hrsJe^r( Z'g I 6ZI uorpnporlul I'9 ) €'8 s z'8 6Zl suraDEd.tftuopnlo^fi v r'8 'lqeN pu? uosuerge"p4uopoeloJ qal oql '8 tzr ??6I I.0l.s tzr u,l^Irrrrs{ ou)lun JspJO 0I'S zzl 196l 'sapoqupue u4snv a?pqleuaolse^flurq €1.6.s 'sepoqupu? u48nv e"pqleuEsn^?Jr{lrureJ L 0zl I86I zl.6.s 'reulu e"pqleuEolee^\s,{Iulec L Ltl 9g6l I I.6's SII 'IJelJ eap4uopoqpuftqruvi(Iu"c {. 9'L 7,L61 01.6.s €II 'quoiasauuloHpu? srxeH eeprluopoql"u8orpl L e€6I 6.6's III snqQuSpSoApue DaUqroT^lrtued g'6 L S ll\au.eep4uopoqleuc dlrued L 60r L.6.s 'sepoqupu? urlsnv eepqleuSolcrla L 90r Ig6I 9.6.9 ,reu.e?prdelolerul?d^lluleg^lrued I SL z0l 5.6.5 .relssegesprqlsuai(lod flruleg L 66 sz6l ,.6.s 'redooJeup4uopornBdsoreld ]{lrurEJ L 86 LL6l E 6.s Ltt 'reddelxeeprTlele{co) {[u]BJ L Ig6I 2.6.5 s ?'r 06 6S6l 'sseHe"p4uopoqleu8oqteds I.6.s 68 'Irzq ?prurpolrBzo^IruBJ eqJ L 916l 6's L8 'uro.4spurla?prTlelopuoc /(Iuleg L 016I s.g.s g tL s8 zL6l'TrelJeuprruoslllg/([rrrEg t.g.s .ur-o.4spur.Ie"pr-rlleuEorlceg I zL €8 016l r{lrureJ €.9.s I8 :relssege"puporuogd /{lrluec q IL 9Z6l z.g.s OL tt6l 'Iqew pue uosu?rge?prqleutorqc {IIu"c I.g's FOslEd '' 8L ,{\ou'eprurporuoFd aql g s 9L 016I'ruo4spur-IsBprluopouod,(lruJel zI. L's SL 016I 'urgrtspur.I*pqtsuEoprdrqu ll' L.s s 8'9 NL 6S6l 'ss?Hosprluoporuo!^J ,tllru?{^lrru?d 0I.L.s 9 ct ,r\eu'eeprurpopald ,qFued 6'L'g 9 ZL tg6l 'slrrsH e"p4uopolsro4lnll g.t.s rruorlstrege"pql"utoceld^lod^llure{ x L9 IL Ig6l ^Irue{ L.L-9 9 69 ts6l 'rellnn pu? relml I e?pnuopoucl.,(Iuleg g L.s 9 /o Ig6I 'redd"Dl eepquopouolsr(IJ{lruleg S-L-S 9 s9 1l\eu'eepqlepoucl/(IIuIug v'L's f. 9'9 t9 6S6l 'sseHe"pqleuEopg ,{IurBJ E L.s rI s'9 z9 9z6l 'r3lss?ge?pquoporuoud {lrrrrBJ z.L's a v9 09 016l 'ur-o.4spur.Ie"puuopolsro lL.s 9 6S 9L6I'I:IZ(, sprluoporuoud^lruru eqJ ,'s r 9 9S 0t6I'rrrojtspur.I e?prluopolepued^lrruBJ I.9.S v 9 ss spquoporepuedeql 9'9 hl SJNAINOC IITA L \,,, CONTENTS lx 55 6.3.3 Siluriancycles r34 fb 6.3.4 Devonian and Carboniferouscycles 135 59 6.3.5 Permian and Triassic cycles r37 60 6.4 Extinction 138 62 6.5 Iterativeevolutionarypatterns 140 63 6.6 Evolutionary trends t42 65 6.6.1 Apparatuselaboration t42 67 6.6.2 Apparatusreduction 143 69 6.6.3 Elaboration of elementsin P positions 144 7l 6.7 Developmentalslralegies 145 72 6.7.1 Recapitulation 145 6.7.2 Paedomorphosrs t46 74 6.8 Summary t46 75 76 78 7. Paleoecologyand paleobiogeography 149 79 8l 7.1 Introduction 149 83 7.2 Mode of life, or habit, of conodonts 149 7.3 Ecologicmodels 6) 150 87 1.3.1 Thedepth-stratificationmodel l5l 89 1.3.2 The lateral-segegationmodel 152 90 7.4 Selectedstudies of conodont ecology 152 9'7 7.4.1 Ordovician paleoecologyof Cincinnati Region t52 98 7.4.2 Mississippianpaleoecology, western United States 157 99 7.4.3 Paleoecologyof Pennsylvanianconodonts 160 t02 7.5 Ecologicgeneralizations 164 r06 7.5.1 Depth as a factor 165 109 7.5.2 Ternperutureas a factor 165 tlt 7.5.3 Nearshoreand ofshore faunas 165 7.5.4 Phyletic changesin ecologicaldistribution l_ - ::l 113 t66 115 7.6 Paleobiogeography 166 |7 7.6.1 Late Cambrian and Ordovician paleobiogeography t67 t20 7.6.2 l-ater Paleozoicand Triassic paleobiogeography 167 122 123 8, The phylum Conodonta 170 t23 8.1 A personalbias 170 t29 8.2 Summary of conodont characters 170 8.3 Conodonts as invertebrates t7l r29 8.3.1 Arthropod and annelid connections t7l r30 8.3.2 Molluscanconnections 172 132 8.3.3 Connectionswith other invertebrates 173 132 8.4 Conodonts as chordates 175 133 8.4.1 The opinions of Pander 175 133 8.4.2 Newberry, Hinde, Huxley and Myxine r75 t34 8.4.3 Macfarlane and the nemertineanconnection 176 s07 XAPUI 68r sueqc e?uerrrqder8uats g xrpuaddv s8l etuopouoJ eql Jo uo4?cgrsselctuputuns v v xpuoddv z8l ro^ug..I g.g I8I seuru{geol?pJoqc ?urlecrpur sernlee3 Jo fu?ruruns 6.t.g 6LI auxftl,tJ1o snteredd?pnAuq eqJ g.r.g suourceds LLI snolgJruoqJ?Jqsr ors eqlJo suorlelsrfualuy 4.7.g LLI slualuoleJo qyro:8 pue uoqrsodruoJ 9.t.9 9LI IBllaletu ,{uoq pseq Jo uorl"laJfuelur pue fus^ocsrq S.r'g 9LI relsssgpu? qslrlnJoJerTeq eqL n.v.g troJ eqJ SJ-NETNOi) The Conodonta 176 t76 t77 t77 t79 181 t82 185 189 lu) sr Jl ? -ro j I SUOIlqer ,i(qroN h or e,ia ',(e^rnslmrEoloaC srourlll ,{q peqsruJnJ,tlpuDl se^ntsod -EIp Jolo3 uro+ apeIII quud 'snpoqwug ^oupoU P SJTUEe_I elrq^\ pue \Jelg Jo seroads_ed^lpel?u8rsap ,{lluacor s?^{ rr qcrgA '(,(punou) snpaultrq snpoqpug slueseJderpue eueluol uI $lcoJ snoJeJruoqffCrando-I t FxuJs .srourll I , il sruoq uro{ sr e8elqueswurouog ur $IcoJ(u"unossrl I) ueru?Al^suusdruo{ ,(sepoql{)snrld,t mqpuSo .gtx ,slueuale |$O-XJ Jq us Jo ed,tloloqsr dol tp atelqruassv luopouocJosaEelqruesie prnlep F suopo r {qr uI 'tuJsard + m uJrq F J{OZOel lfuL\los le lxxroJ Jql q leql lnq cfed srql D :{IO lsEAI lE-fueduroJ slrsso_I qer naql :qr:roru b ;mssed ?rI _to fg G rGololq hs Eruoq |D 'ulued r|e F:ujeal ppol-t ou1 r -iqdB-6'u Fdar rsot! &1u.uo5 x sdno-6 IP|:-j llJto lEseq lual lsqf, ale.$ t D Jrqde6 3122\ $.el e lEi sFsso_I FB PaESU lsso_loDlur lla-rur luJn FisE lsour P IJOJ Juu r ls.n! 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    MILWAUKEE PUBLIC MUSEUM Contributions . In BIOLOGY and GEOLOGY Number 83 March 1,1992 A Compendium of Fossil Marine Animal Families 2nd edition J. John Sepkoski, Jr. MILWAUKEE PUBLIC MUSEUM Contributions . In BIOLOGY and GEOLOGY Number 83 March 1,1992 A Compendium of Fossil Marine Animal Families 2nd edition J. John Sepkoski, Jr. Department of the Geophysical Sciences University of Chicago Chicago, Illinois 60637 Milwaukee Public Museum Contributions in Biology and Geology Rodney Watkins, Editor (Reviewer for this paper was P.M. Sheehan) This publication is priced at $25.00 and may be obtained by writing to the Museum Gift Shop, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, WI 53233. Orders must also include $3.00 for shipping and handling ($4.00 for foreign destinations) and must be accompanied by money order or check drawn on U.S. bank. Money orders or checks should be made payable to the Milwaukee Public Museum. Wisconsin residents please add 5% sales tax. In addition, a diskette in ASCII format (DOS) containing the data in this publication is priced at $25.00. Diskettes should be ordered from the Geology Section, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, WI 53233. Specify 3Y. inch or 5Y. inch diskette size when ordering. Checks or money orders for diskettes should be made payable to "GeologySection, Milwaukee Public Museum," and fees for shipping and handling included as stated above. Profits support the research effort of the GeologySection. ISBN 0-89326-168-8 ©1992Milwaukee Public Museum Sponsored by Milwaukee County Contents Abstract ....... 1 Introduction.. ... 2 Stratigraphic codes. 8 The Compendium 14 Actinopoda.
  • Dhanda, R., Murdock, DJE, Repetski, JE, Donoghue, PCJ, & Smith, MP

    Dhanda, R., Murdock, DJE, Repetski, JE, Donoghue, PCJ, & Smith, MP

    Dhanda, R., Murdock, D. J. E., Repetski, J. E., Donoghue, P. C. J., & Smith, M. P. (2019). The apparatus composition and architecture of Erismodus quadridactylus and the implications for element homology in prioniodinin conodonts. Papers in Palaeontology, 5(4), 657-677. https://doi.org/10.1002/spp2.1257 Peer reviewed version Link to published version (if available): 10.1002/spp2.1257 Link to publication record in Explore Bristol Research PDF-document This is the author accepted manuscript (AAM). The final published version (version of record) is available online via Wiley at https://onlinelibrary.wiley.com/doi/full/10.1002/spp2.1257 . Please refer to any applicable terms of use of the publisher. University of Bristol - Explore Bristol Research General rights This document is made available in accordance with publisher policies. Please cite only the published version using the reference above. Full terms of use are available: http://www.bristol.ac.uk/red/research-policy/pure/user-guides/ebr-terms/ THE APPARATUS COMPOSITION AND ARCHITECTURE OF ERISMODUS QUADRIDACTYLUS AND THE IMPLICATIONS FOR ELEMENT HOMOLOGY IN PRIONIODININ CONODONTS By ROSIE DHANDA1, DUNCAN J. E. MURDOCK2, JOHN E. REPETSKI3 and PHILIP C. J. DONOGHUE4 and M. PAUL SMITH2 * 1School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK; [email protected] 2 Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW, UK; [email protected], [email protected] 3 U.S. Geological Survey, MS 926A
  • Conodonts (Vertebrata)

    Conodonts (Vertebrata)

    Journal of Systematic Palaeontology 6 (2): 119–153 Issued 23 May 2008 doi:10.1017/S1477201907002234 Printed in the United Kingdom C The Natural History Museum The interrelationships of ‘complex’ conodonts (Vertebrata) Philip C. J. Donoghue Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queen’s Road, Bristol BS8 1RJ, UK Mark A. Purnell Department of Geology, University of Leicester, University Road, Leicester LE1 7RH, UK Richard J. Aldridge Department of Geology, University of Leicester, University Road, Leicester LE1 7RH, UK Shunxin Zhang Canada – Nunavut Geoscience Office, 626 Tumit Plaza, Suite 202, PO Box 2319, Iqaluit, Nunavut, Canada X0A 0H0 SYNOPSIS Little attention has been paid to the suprageneric classification for conodonts and ex- isting schemes have been formulated without attention to homology, diagnosis and definition. We propose that cladistics provides an appropriate methodology to test existing schemes of classification and in which to explore the evolutionary relationships of conodonts. The development of a multi- element taxonomy and a concept of homology based upon the position, not morphology, of elements within the apparatus provide the ideal foundation for the application of cladistics to conodonts. In an attempt to unravel the evolutionary relationships between ‘complex’ conodonts (prioniodontids and derivative lineages) we have compiled a data matrix based upon 95 characters and 61 representative taxa. The dataset was analysed using parsimony and the resulting hypotheses were assessed using a number of measures of support. These included bootstrap, Bremer Support and double-decay; we also compared levels of homoplasy to those expected given the size of the dataset and to those expected in a random dataset.
  • ABSTRACT BOOK a Cura Della Società Geologica Italiana

    ABSTRACT BOOK a Cura Della Società Geologica Italiana

    https://doi.org/10.3301/ABSGI.2019.04 Milano, 2-5 July 2019 ABSTRACT BOOK a cura della Società Geologica Italiana 3rd International Congress on Stratigraphy GENERAL CHAIRS Marco Balini, Università di Milano, Italy Elisabetta Erba, Università di Milano, Italy - past President Società Geologica Italiana 2015-2017 SCIENTIFIC COMMITTEE Adele Bertini, Peter Brack, William Cavazza, Mauro Coltorti, Piero Di Stefano, Annalisa Ferretti, Stanley C. Finney, Fabio Florindo, Fabrizio Galluzzo, Piero Gianolla, David A.T. Harper, Martin J. Head, Thijs van Kolfschoten, Maria Marino, Simonetta Monechi, Giovanni Monegato, Maria Rose Petrizzo, Claudia Principe, Isabella Raffi, Lorenzo Rook ORGANIZING COMMITTEE The Organizing Committee is composed by members of the Department of Earth Sciences “Ardito Desio” and of the Società Geologica Italiana Lucia Angiolini, Cinzia Bottini, Bernardo Carmina, Domenico Cosentino, Fabrizio Felletti, Daniela Germani, Fabio M. Petti, Alessandro Zuccari FIELD TRIP COMMITTEE Fabrizio Berra, Mattia Marini, Maria Letizia Pampaloni, Marcello Tropeano ABSTRACT BOOK EDITORS Fabio M. Petti, Giulia Innamorati, Bernardo Carmina, Daniela Germani Papers, data, figures, maps and any other material published are covered by the copyright own by the Società Geologica Italiana. DISCLAIMER: The Società Geologica Italiana, the Editors are not responsible for the ideas, opinions, and contents of the papers published; the authors of each paper are responsible for the ideas opinions and con- tents published. La Società Geologica Italiana, i curatori scientifici non sono responsabili delle opinioni espresse e delle affermazioni pubblicate negli articoli: l’autore/i è/sono il/i solo/i responsabile/i. © Società Geologica Italiana, Roma 2019 STRATI 2019 ABSTRACT INDEX ST1.1 History of Stratigraphy in Italian environments (17th – 20th centuries) ........................................