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Morphological Variation in the Hadrosauroid Dentary Morfologisk Variation I Det Hadrosauroida Dentärbenet

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Morphological Variation in the Hadrosauroid Dentary Morfologisk Variation I Det Hadrosauroida Dentärbenet Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 398 Morphological Variation in the Hadrosauroid Dentary Morfologisk variation i det hadrosauroida dentärbenet D. Fredrik K. Söderblom INSTITUTIONEN FÖR GEOVETENSKAPER DEPARTMENT OF EARTH SCIENCES Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 398 Morphological Variation in the Hadrosauroid Dentary Morfologisk variation i det hadrosauroida dentärbenet D. Fredrik K. Söderblom ISSN 1650-6553 Copyright © D. Fredrik K. Söderblom Published at Department of Earth Sciences, Uppsala University (www.geo.uu.se), Uppsala, 2017 Abstract Morphological Variation in the Hadrosauroid Dentary D. Fredrik K. Söderblom The near global success reached by hadrosaurid dinosaurs during the Cretaceous has been attributed to their ability to masticate (chew). This behavior is more commonly recognized as a mammalian adaptation and, as a result, its occurrence in a non-mammalian lineage should be accompanied with several evolutionary modifications associated with food collection and processing. The current study investigates morphological variation in a specific cranial complex, the dentary, a major element of the hadrosauroid lower jaw. 89 dentaries were subjected to morphometric and statistical analyses to investigate the clade’s taxonomic-, ontogenetic-, and individual variation in dentary morphology. Results indicate that food collection and processing became more efficient in saurolophid hadrosaurids through a complex pattern of evolutionary and growth-related changes. The diastema (space separating the beak from the dental battery) grew longer relative to dentary length, specializing food collection anteriorly and food processing posteriorly. The diastema became ventrally directed, hinting at adaptations to low-level grazing, especially in younger individuals. The coronoid process became anteriorly directed, and was relatively more elongate, resulting in increased moment arm length, with muscles being re-directed to pull the jaw more posteriorly, and mechanical advantage increasing. Although all hadrosauroid groups went through relative dental battery elongation during growth, by incorporating more teeth into each row, the dental battery became deeper in saurolophids. Previous research supports the interpretation that this is the result of more tooth rows being stacked vertically, allowing the dental battery to work as a shock-absorber during mastication, and allowed teeth to be replaced without interruption to food consumption. The increased anterior inclination of the coronoid process, and relative elongation of the diastema in saurolophids are herein suggested to have evolved through hypermorphosis, a version of peramorphosis where the growth trajectory in the descendant taxon extends beyond the ancestral state, whereas the relative elongation of the coronoid process, the relative deepening of the dental battery, and the increased ventral deflection of the diastema are the result of a novel juvenile condition. Keywords: Hadrosauroidea, dentary, morphometric, heterochrony, peramorphosis, mastication Degree Project E1 in Earth Science, 1GV025, 30 credits Supervisor: Nicolás E. Campione Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala (www.geo.uu.se) ISSN 1650-6553Examensarbete vid Institutionen för geovetenskaper, No. 398, 2017 The whole document is available at www.diva-portal.org Populärvetenskaplig sammanfattning Morfologisk variation i det hadrosauroida dentärbenet D. Fredrik K. Söderblom Hadrosauroider, även kallade ank-näbbs dinosaurier, var vanligt förekommande och nästan globalt distribuerade växtätare som levde under krita-perioden. Forskning har länge tytt på att deras framgång berott på deras förmåga att processera föda genom att tugga, en förmåga som länge ansågs exklusiv till däggdjur. Denna förmåga förbättrades sedan genom morfologiska förändringar som dök upp i de saurolophida hadrosauroidera. Dessa förändringar ledde till både effektivare födoinsamling och bearbetning. En serie analyser utfördes på 89 dentärben (ett av benen i underkäken) för att testa hur dessas form varierade inom Hadrosauroidea, beroende på gruppers släktskap, ålderstadie, samt individuell variation. Resultaten tyder på att diastemat, området som i hadrosauroider separerar predentärbenet från dental batteriet (liknande hur fram- och kindtänderna är separerade hos hästar) blev längre i saurolophider. Detta tillät ökad specialisering av predentärbenet till insamling av mat och i dental batteriet (rader av tätt sittande tänder placerade vertikalt på varandra) till att mala ned mat. Diastemats ventrala böjning avtog i alla grupper allt eftersom individen åldrades, men var i helhet mer böjt i saurolophider. Detta tolkas som möjlig anpassning till betande på lågt växande vegetation, som idag ses i vissa hovdjur (Ungulata). Koronoid processen, en vertikalt utstickande del av dentärbenet som är placerad långt bak, lutades framåt och förlängdes i saurolophider. Denna förändring riktade muskler till att dra käken mer bakåt, samt ökade hävstångsverkan, vilket ökade saurolophiders bitkraft. Alla hadrosauroider gick igenom förlängning av dental batteriet genom tillägg av tänder, vilket ökade tuggytans area. Saurolophiders dental batterier var djupare än sina föregångares, vilket har dokumenterats i litteratur som ännu fler vertikalt staplade rader. Fler tandrader betydde att stötar från starkare bitkraft kunde absorberas över en större yta. Dessutom så kunde tandrader därför bytas utan dröjsmål när den översta raden nötts ned, vilket tillät oavbruten födokonsumtion. Anledningen till den ökade lutningen i koronoid processen, och förlängningen av diastemat tros vara hypermorfos, vilket är när tillväxten av något som slutade växa i förfäderna istället fortsätter växa i ättlingarna. Den ökade längden på koronoid processen, det djupare dental batteriet, och det mer ventralt böjda diastemat i saurolophider uppstod p.g.a. att ett nytt juvenilt tillstånd utvecklades. Nyckelord: Hadrosauroidea, dentärben, morfometrisk, heterokroni, peramorfos, tugga Examensarbete E1 i geovetenskap, 1GV025, 30 hp Handledare: Nicolás 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 398, 2017 Hela publikationen finns tillgänglig på www.diva-portal.org Table of Contents 1 Introduction .......................................................................................................................................1 2 Aims ....................................................................................................................................................2 3 Background ........................................................................................................................................4 3.1 Cretaceous climate and environment ..........................................................................................4 3.2 Taxonomic conventions and the dentary’s use in phylogenetics ................................................4 3.3 Evolution of the feeding apparatus and jaw mechanics in hadrosauroids ...................................7 3.4 Examples of previous research on morphological variation in hadrosauroid dentaries ..............9 3.4.1 Taxonomic variation ...........................................................................................................9 3.4.2 Dentary growth patterns ....................................................................................................10 3.4.3 Individual variation, taphonomic-, and pathologic alteration ...........................................11 4 Methods ............................................................................................................................................13 4.1 Data collection...........................................................................................................................13 4.2 Digitization ................................................................................................................................13 4.3 Generalized procrustes analysis and semilandmarks sliding .....................................................14 4.4 Principal component analysis, morphospace diagram and density plots...................................15 4.5 Normality tests and principal component group separation ......................................................15 4.6 Bivariate allometry plots and confidence intervals ...................................................................16 5 Results ..............................................................................................................................................17 5.1 Morphospace diagram and density plots ...................................................................................17 5.1.1 Principal component 1 .......................................................................................................17 5.1.2 Principal component 2 .......................................................................................................19 5.2 Normality tests ..........................................................................................................................21 5.3 Tukey’s honest significant difference test .................................................................................22
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