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Disparity of Early Cretaceous Lamniformes Sharks Disparitet I Lamniformes Hajar Från Tidig Krita

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Disparity of Early Cretaceous Lamniformes Sharks Disparitet I Lamniformes Hajar Från Tidig Krita Independent Project at the Department of Earth Sciences Självständigt arbete vid Institutionen för geovetenskaper 2015: 7 Disparity of Early Cretaceous Lamniformes Sharks Disparitet i Lamniformes Hajar från Tidig Krita Fredrik Söderblom DEPARTMENT OF EARTH SCIENCES INSTITUTIONEN FÖR GEOVETENSKAPER Independent Project at the Department of Earth Sciences Självständigt arbete vid Institutionen för geovetenskaper 2015: 7 Disparity of Early Cretaceous Lamniformes Sharks Disparitet i Lamniformes Hajar från Tidig Krita Fredrik Söderblom Copyright © Fredrik Söderblom and the Department of Earth Sciences, Uppsala University Published at Department of Earth Sciences, Uppsala University (www.geo.uu.se), Uppsala, 2015 Sammanfattning Disparitet i Lamniformes Hajar från Tidig Krita Fredrik Söderblom Morfologisk disparitet är ett mått på hur stor utsträckningen av morfologisk variation är. Detta mått räknas ut genom att jämföra landmärken utplacerade på bilder av föremål som ska undersökas. I detta projekt undersöktes den morfologiska dispariteten hos tänder från håbrandsartade hajar (Lamniformes) under tidig krita. Att just deras tänder undersöktes beror på att den större delen av hajars skelett är gjort av brosk vilket lätt bryts ned efter djuret avlidit. Deras tänder är dock gjorda av ben vilket har lättare att bli bevarat som fossil. Utöver detta så kan formen på tänder beskriva djurs födoval och levnadssätt. Gruppens tänder undersöktes därför även för att belysa eventuella förändringar i diet och ekologi under tidig krita. Resultatet av denna analys visar på en expansion av tandform under denna period från långa och smala tänder under Barremium till en större variation under Albium där även mer triangelformade och robusta tänder dyker upp. Detta har tolkats som en adaptiv artbildningsperiod för gruppen då både nya byten (t.ex. teleostfiskar och havs- sköldpaddor) diversifierade och uppkom samtidigt som vissa marina predatorer (ichthyosaurer och plesiosaurer) minskade i antal under denna tidsperiod. Detta ändrade troligen de selektiva trycken på håbrandsartade hajars tandmorfologi samt lämnade ekologiska nischer öppna som dessa kunde anpassa sig till vilket i sin tur ledde till expansioner i morfologisk disparitet, diet och ekologi. Nyckelord: Lamniformes, disparitet, tidig krita, morfologi, morfometri Självständigt arbete i geovetenskap, 1GV029, 15 hp, 2015 Handledare: Nicolàs Campione Biträdande handledare: Benjamin Kear Institutionen för geovetenskaper, Uppsala universitet, Villavägen 16, 752 36 Uppsala (www.geo.uu.se) Hela publikationen finns tillgänglig på www.diva-portal.org Abstract Disparity of Early Cretaceous Lamniformes sharks Fredrik Söderblom The geological range of lamniform sharks stretches from present day species such as Carcharodon carcharias (great white shark) back to the at the moment oldest undoubted fossil finds during the Early Cretaceous. In this paper a geometric morphometric analysis was performed on images of Early Cretaceous lamniform teeth collected from published literature in order to examine the change in disparity (range of morphological variation within a group) throughout the time period. Due to limited availability of published material and time constraints only the Barremian and Albian ages were investigated. The Barremian exhibited tall and narrow tooth morphologies while the Albian showed a wide range of morphological variation including more robust, wide and sometimes triangular shapes but also displayed further specialization of the tall and narrow forms. This change is likely indicative of a dietary and ecological expansion from only eating for example small fish and soft- bodied creatures to a wide range of prey for the group, including larger and more robust animals such as marine turtles and large bony fish. This in combination with the decline of some marine predators as well as the diversification of possible prey is interpreted as that an adaptive radiation of the Lamniformes could have taken place during the latter half of the Early Cretaceous. Key words: Lamniformes, disparity, Early Cretaceous, morphology, morphometric Independent Project in Earth Science, 1GV029, 15 credits, 2015 Supervisor: Nicolàs Campione Co-Supervisor: Benjamin Kear Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala (www.geo.uu.se) The whole document is available at www.diva-portal.org Table of contents 1. Introduction 1 2. Materials and methods 2 3. Results 3 4. Discussion 5 4.1 Discussion of the results 5 4.2 Dental adaptations and habitats 6 4.2.1 The Barremian 6 4.2.2 The Albian 8 4.3 On changes in disparity, diversity, diet and feeding strategies 10 5. Conclusions 13 6. Acknowledgements 14 7. References 14 7.1 Printed refereces 14 7.2 Internet references 17 8. Appendix 17 1. Introduction Chondrichthyes is the name of a group including the clades Holocephali (chimaeras) as well as Elasmobranchii (sharks and rays) (Miller et al., 2003; Hickman et al., 2011). Fossil scales prove chondrichthyans to possibly have existed since the Late Ordovician (Janvier, P., 1996 & Turner, S. in Miller et al., 2003). They are fish with a skeleton made of cartilage, a feature that developed as they evolved out of their ancestors with skeletons made of bone (Hickman et al., 2011). The fact that their teeth are better mineralized than the rest of their skeleton is the reason that teeth are one of the most common kind of fossil found belonging to them since cartilage is less likely to survive the fossilization process (Shimada, 2005, 2007; Whitenack & Gottfried, 2010). Sharks that are a part of them have been determined to have existed during the Early Devonian on the basis of fossil teeth (Miller et al., 2003) and even back to the Lower Siluran on the basis of simple placoid scales (Karatajūté- Talimaa, V., 1973, 1992 in Sansom et al., 1996). The clade Lamniformes is an order of sharks that is present on Earth today and includes the great white shark (Carcharodon carcharias) (Cappetta, 2012). The order’s origin was however during the Early Cretaceous with one of the earliest undoubted fossil finds belonging to the early part of the Late Valanginian (Rees, 2005). Although the possibility of an origin during the Jurassic has also been proposed (Underwood, 2006). The Cretaceous is a time period that spans from 145-66 million years ago (Ma) (Cohen et al., 2015). The period is divided into the Early Cretaceous (145- 100.5 Ma) and the Late Cretaceous (100.5-66 Ma) (Cohen et al., 2015). The Early Cretaceous was the focus of this study. It can be further subdivided into the Berriasian (145-139.8 Ma), the Valanginian (139.8-132.9 Ma), the Hauterivian (132.9- 129.4 Ma), the Barremian (129.4-125 Ma), the Aptian (125-113 Ma) and the Albian (113-100,5 Ma) (Cohen et al., 2015). During the beginning of the Early Cretaceous the global climate became more arid than before (Benson & Druckenmiller, 2014) (evidenced in part by clays having an increased proportion of the mineral smectite (Weissert & Channell, 1989; Hallam et al., 1991)), the ocean surface became more oligotrophic (Danelian & Johnson, 2001; Tremolada et al., 2006 in Benson & Druckenmiller, 2014) and temperatures rose to eventually reach a high approximately 100 million years ago (Gould et al., 2001). Radiations of several different organisms took place in the Cretaceous and many of them originated during the Early Cretaceous (Sadava et al., 2011). The first angiosperms (flowering plants) appeared during the Early Cretaceous (Gould et al., 2001; Sadava et al., 2011), triggering a radiation of insects such as butterflies, bees, moths, as well as ants (Gould et al., 2001). The first snakes arose during the Early Cretaceous (Benton, 2005; Sadava et al., 2011). Other organisms such as gymnosperms, teleost fishes, sharks, plesiosaurs, belemnites, ammonites (Gould et al., 2001), dinosaurs (Gould et al., 2001; Sadava et al., 2011), pterosaurs (Gould et al., 2001) (pterodactyloid pterosaurs during the Early Cretaceous (Benson & Druckenmiller, 2014)) and turtles (Early Cretaceous) (Scheyer et al., 2014) would diversify during the Cretaceous. Large predators such as mosasaurs also appeared in the Cretaceous as well as the first true lobsters and crabs (Gould et al., 2001). With this paper a dataset connected to images of Early Cretaceous Lamniformes shark teeth will be presented (see appendix). A geometric morphometric analysis of the teeth in these images was also performed. The results of this analysis will be presented as diagrams. A morphospace diagram showcasing 1 the measure of morphological variation between specimen in a mathematical space and a disparity through time diagram showing the disparity (the range of morphological variation within a group (Briggs & Crowther, 2001)) of lamniform teeth during the different ages of the Early Cretaceous. A t-test was also performed to account for the significance of the sampled specimen in question between the different time bins. The hypothesis being tested is that since a sizeable amount of radiations seem to have taken place at the time investigated in this paper, the results of the analysis might show a high amount disparity and possibly an increase in morphological variation during the aforementioned time period. 2. Materials and methods A total of 146 images of Lamniformes teeth specimen were located using published literature available in both digital and physical formats and compiled into a dataset (see table 1 in appendix). The preferred view in the images was labial (seen from the outside of the mouth),
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