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Shark) Dental Morphology During the Early Mesozoic Dynamik Av Selachian (Haj) Tandmorfologi Under Tidig Mesozoisk

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Shark) Dental Morphology During the Early Mesozoic Dynamik Av Selachian (Haj) Tandmorfologi Under Tidig Mesozoisk Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 399 Dynamics of Selachian (Shark) Dental Morphology During the Early Mesozoic Dynamik av Selachian (haj) tandmorfologi under Tidig Mesozoisk Alexander Paxinos INSTITUTIONEN FÖR GEOVETENSKAPER DEPARTMENT OF EARTH SCIENCES Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 399 Dynamics of Selachian (Shark) Dental Morphology During the Early Mesozoic Dynamik av Selachian (haj) tandmorfologi under Tidig Mesozoisk Alexander Paxinos ISSN 1650-6553 Copyright © Alexander Paxinos Published at Department of Earth Sciences, Uppsala University (www.geo.uu.se), Uppsala, 2017 Abstract Dynamics of Selachian (Shark) Dental Morphology During the Early Mesozoic Alexander Paxinos The ancestors of all modern day sharks and rays (Neoselachii) may have appeared during the Late Palaeozoic, but their major diversification happened sometime during the Early Mesozoic. Taxonomic evidence places the first neoselachian diversification in the Early Jurassic. Taxonomic diversity analyses, however, are often affected by incompleteness of the fossil record and sampling biases. On the other hand, the range of morphological variation (disparity) offers a different perspective for studying evolutionary patterns across time. Disparity analyses are much more resilient to sampling biases than diversity analyses, and disparity has the potential to provide a more ecologically-relevant context. This analysis focuses on the morphology of selachian teeth. Selachii include all neoselachian and stem-neoselachian sharks, but not batoids. In order to primarily test the hypothesis that the first selachian radiation took place in the Late Triassic, not the Early Jurassic, and secondarily to depict the dental evolution of sharks across time, 424 selachian teeth ranging from the Middle Triassic to the Late Jurassic were quantified, using a landmark and semi-landmark geometric morphometric method, in order to analyze patterns of selachian tooth disparity and morphospace occupation across the Early Mesozoic. The results of the analysis indicate a two-pulse radiation of the selachians. The first radiation took place in the Rhaetian (Late Triassic). The Rhaetian Transgression saw the rise of a large, shallow epicontinental sea that covered the area which is now Western Europe. The transgressive event opened new niches for the selachians, as indicated by the appearance of almost all known shark dentition types during the Rhaetian. The second radiation spans the Callovian – Oxfordian – Kimmeridgian interval. Transgressive events during the Callovian and the Oxfordian introduced new cephalopod and actinopterygian faunas to the Central European Basin, thus playing an important role in selachian dental disparity and morphospace patterns. The main drive in shark evolution across the Early Mesozoic seems to have been the breakup of Pangaea. Transgressive/regressive events tied to tectonic activity affect tooth disparity and, indirectly, influence shark dentition patterns, by directly affecting the diversity of other faunas which, in turn, are preyed upon by selachians of the Early Mesozoic. Keywords: Early Mesozoic, Selachii, morphological disparity, geometric morphometrics, early radiation Degree Project E1 in Earth Science, 1GV025, 30 credits Supervisor: Nicolàs E. Campione Department of Earth Sciences, Uppsala University, Villavägen 16, 752 36 Uppsala (www.geo.uu.se) ISSN 1650-6553, Examensarbete vid Institutionen för geovetenskaper, No. 399, 2017 The whole document is available at www.diva-portal.org Populärvetenskaplig sammanfattning Dynamik av Selachian (haj) tandmorfologi under Tidig Mesozoisk Alexander Paxinos Förfäderna i alla moderna hajar och strålar (Neoselachii) kan ha uppträtt under den sena palæozoiska, men deras stora diversifiering hände någon gång under den tidiga mesozoiska. Nyliga diversifieringsstudier baserade på antalet taxa, placerar den första neoselachiska diversifieringen för 199-190 miljoner år sedan, i tidiga jurassisk. De taxonomiska diversifiering analyser kan ofta bli påverkade av ofullständigheter i den fossila posten och även av den provtagnings tekniken. Längten av de morfologiska variationen (disparitet) ger ett annat perspektiv för att kunna studera evolutionära mönster över tiden. Disparitetsanalyser är mindre påverkade av provtagnings problem och ibland av de fåtaliga prov siffrorna. Studien av morfologiska mönster av biologiska strukturer (såsom tänder) kan markera beteendemönster relaterade till ekologin hos en grupp av organismer (såsom hajar), till och med villebråd val och föredragna livsmiljöer. Den här analysen fokuserar på morfologin av selachian tänder. I Selachimorpha tillhör alla neoselakiska hajar och deras förfäder, men inte strålar eller rockor. En geometrisk morfometrisk metod som använder två dimentionella, semi landmärken är implimenterad i den här studien för att testa hypotesen att den första selachiska utstrålningen ägde rum i den sena triassiska ~ 10 miljoner år tidigare än diversitets analyser hade uppskattat. Morfologin av 424 hajar tänder som tillhör 50 genera och 71 arter och spänner ifrån den mellersta triasisk till den sena jurassisk var kvantifierad genom användning av kurvor. Resultaten av analysen indikerar att selachierna diversifierades två gånger under den tidiga mesozoiska. Den första utstrålningen ägde rum i den Rhaetian (sen triassisk).Den Rhaetian Transgression såg uppkomsten av ett stort, grundt epikontinentalt hav som täckte en område som idag är Västeuropa. Den transgressiva händelsen öppnade nya ekologiska möjligheter för selachierna, vilket framgår av utseendet på nästan alla kända haj-dentitionstyper under Rhaetian. Den andra utstrålningen sträcker sig över Callovian-Oxfordian-Kimmeridgian-intervallet. Transgressiva händelser under Callovian och Oxfordian orsakade havsnivån att stiga och introducerade nya cefalopoder och aktinopterygiska faunor till de epikontinentala haven av Centraleuropa och kunde ha spelat därmed en viktig roll i selakiska tands disparitet och morfologisk mönster. Huvudfaktoren i hajutveckling över den tidiga mesozoiska kunde ha varit uppdelningen av Pangaea. Havsnivån fluktuationer orsakade av tektonisk aktivitet kopplad till Pangaeans uppdelningen kunde ha påverkat tändernas disparitet och indirekt orsakat förändringar i haj-dentitioner, genom att direkt påverka diversiteten hos andra faunor, som i sin tur är jagade av selachier från den tidiga mesozoiska. Nyckelord: Tidig Mesozoisk, Selachimorpha, morfologisk disparitet, geometrisk morfometri, tidig strålning Examensarbete E1 i geovetenskap, 1GV025, 30 hp Handledare: Nicolàs E. Campione Institutionen för geovetenskaper, Uppsala Universitet, Villavägen 16, 752 36 Uppsala (www.geo.uu.se) ISSN 1650-6553, Examensarbete vid Institutionen för geovetenskaper, Nr 399, 2017 Hela publikationen finns tillgänglig på www.diva-portal.org Table of Contents 1 Introduction .........................................................................................................................................1 2 Background ..........................................................................................................................................3 2.1 Diversity patterns ...........................................................................................................................3 2.2 Taxonomic richness vs morphological disparity ............................................................................3 2.3 Shark teeth ......................................................................................................................................4 3 Objectives & Hypotheses ....................................................................................................................6 4 Methods ................................................................................................................................................7 4.1 Data collection ................................................................................................................................7 4.2 Cataloguing & stratigraphy ............................................................................................................8 4.3 Geometric morphometrics ..............................................................................................................9 4.4 Digitization & resampling ..............................................................................................................9 4.5 Analysis ........................................................................................................................................11 4.6 Morphological disparity ...............................................................................................................12 4.7 Sample bias ...................................................................................................................................12 4.8 Clade-specific disparity analysis ..................................................................................................12 5 Results ................................................................................................................................................14 5.1 Disparity patterns .........................................................................................................................14 5.2 Morphospace patterns ...................................................................................................................16
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