University of Alberta Paleobiology of Canadian Cretaceous amber: an exceptional record of Late Cretaceous Hymenoptera, with contributions to additional taxa and the study of amber by Ryan Christopher McKellar A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Earth and Atmospheric Sciences ©Ryan Christopher McKellar Fall 2011 Edmonton, Alberta Permission is hereby granted to the University of Alberta Libraries to reproduce single copies of this thesis and to lend or sell such copies for private, scholarly or scientific research purposes only. Where the thesis is converted to, or otherwise made available in digital form, the University of Alberta will advise potential users of the thesis of these terms. 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While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. ••I Canada ABSTRACT Canadian amber provides a glimpse of a Late Cretaceous (Campanian) ecosystem preserved in exquisite detail. Here we document one of the best- preserved assemblages of Hymenoptera (ants, bees, and wasps) in the Mesozoic. Contributions are also made to the study of a broader range of taxa and the development of amber research. Amber offers the best fossil record for diminutive terrestrial organisms with a low preservation potential, such as micro-hymenopterans. Canadian amber preserves a broad range of hymenopteran taxa, but is strongly biased towards smaller taxa, so members of 'Parasitica' (a grade of parasitoid hymenopterans) are most thoroughly represented. To date, 30 families, 38 genera and 70 species of Hymenoptera have been identified in the assemblage: 36 of these species, six of the genera, and two of the families stem from the body of work summarized in this dissertation. Herein, Canadian amber specimens are contrasted against those recovered from other Cretaceous amber deposits, and large-scale palaeobiogeographic and stratigraphic range patterns are discussed. New taxa are documented within Neuroptera (lacewings) and Hemiptera (true bugs), as these specimens constitute important records for their respective families (Rhachiberothidae and Microphysidae). The most abundant and diverse assemblage of feathers and putative protofeathers yet to be recovered from Mesozoic amber is also described. Careful screening of Canadian amber collections for inclusions as small as micro-hymenopterans has yielded a wide range of inclusions of high scientific value. Contributions to the study of these specimens will promote additional research into a somewhat overlooked palaeontological resource. Finally, the versatility of stable isotope analyses in the study of amber is demonstrated. This technique is used to explore the source of Canadian amber, and the role of insect attacks in the formation of amber deposits. This new approach to the study of amber provides a more comprehensive account of conditions in the amber-producing forest, and allows palaeoecological hypotheses to be tested through modern analogues. ACKNOWLEDGEMENTS This work would not have been possible without the support, collaboration and mentorship of many people. I am greatly indebted to both of my supervisors, Brian Chatterton and Alex Wolfe, for their support, guidance and encouragement in all facets of my research. I would like to thank Allan Ashworth, Michael Caldwell, Karlis Muehlenbachs, and Felix Sperling for acting as my committee members and examiners - their time, expertise and guidance has been invaluable. My family has been extremely supportive throughout all of my studies, and deserves a great deal of credit here. Collaborative research with Michael Engel (University of Kansas) and Jaime Ortega-Bianco (Universitat de Barcelona) in particular has expanded my understanding and experience with palaeoentomology. Work with Karlis Muehlenbachs and Ralf Tappert has greatly improved my grasp of stable isotope geochemistry and FTIR analyses, while research with Stacey Gibb has contributed to my understanding of ichnology and invertebrate palaeontology. Numerous other workers, particularly those that have acted as collaborators, provided invitations to contribute works, or edited and reviewed the papers encompassed by this dissertation have also provided opportunity for growth. As with most palaeontological projects, specimens have been essential in this study. I am extremely grateful to those that have donated specimens for my research, such as the Leuck family; or have provided access to museum collections, such as Andrew Bennett, Jim Troubridge, and Robert Skidmore (Canadian National Collection of Insects and Arthropods); James Gardner and Brandon Strilisky (Royal Tyrrell Museum of Palaeontology); Philip Perkins (Harvard Museum of Comparative Zoology); Danny Shpeley and Felix Sperling (University of Alberta Strickland Entomological Museum); Andrew Locock (University of Alberta Palaeontology Museum); Nicola Howard (University of Alberta Laboratory for Vertebrate Palaeontology); Janet Waddington and Antonia Guidotti (Royal Ontario Museum); and Jocelyn Hudon (Royal Alberta Museum). Numerous friends and colleagues have also shared their experience, permitted the use of their equipment, or have assisted with fieldwork, including: Kevin Aulenback, Bruce Archibald, Amber Garrett, Will Hobbs, Shane and Vicki Leuck, Svetlana Kuzmina, Lindsey Leighton, Allan Lindoe, Sergei Matveev, Valerie McKellar, Darrin Molinaro, Heather Proctor, Thomas Stachel, Ralf and Michelle Tappert, and Graham Young. Funding was provided through NSERC (PGS D-3) and Alberta Ingenuity Fund (AIF Studentship) support, as well as NSERC Discovery Grants (to B.D.E. Chatterton and A.P. Wolfe). TABLE OF CONTENTS ABSTRACT i ACKNOWLEDGEMENTS iii LIST OF TABLES viii LIST OF FIGURES ix LIST OF SYMBOLS, NOMENCLATURE, OR ABBREVIATIONS xii CHAPTER 1:AGENERAL INTRODUCTION TO CANADIAN AMBER 1 OBJECTIVES OF STUDY AND PROGRESSION OF PAPERS 1 INTRODUCTION 2 GEOLOGICAL SETTING 4 AMBER COLLECTION 5 PALAEOHABITAT 7 THE CANADIAN AMBER TREE 7 AGE OF CANADIAN AMBER 11 PHYSICAL AND CHEMICAL PROPERTIES 12 THE DIVERSITY OF CANADIAN AMBER INCLUSIONS 13 REFERENCES 19 CHAPTER 2: CORRELATION OF GRASSY LAKE AND CEDAR LAKE AMBERS USING INFRARED SPECTROSCOPY, STABLE ISOTOPES, AND PALAEOENTOMOLOGY 26 INTRODUCTION 26 Historical background 26 Cedar Lake amber 26 Grassy Lake amber 27 Stratigraphic constraints 28 Objectives of study 29 MATERIALS AND METHODS 30 Materials 30 FTIR spectroscopy 30 Stable isotopes 31 Palaeoentomology 32 RESULTS 32 FTIR spectra 40 Stable isotopes 42 Amber morphology and physical properties 44 Palaeoentomology 46 DISCUSSION 47 Amber spectroscopy, weathering, and botanical source 47 Inferences from the stable isotopic composition of amber 49 Palaeoentomological and stratigraphic considerations 51 CONCLUSIONS 53 REFERENCES 55 CHAPTER 3: HYMENOPTERA IN CANADIAN CRETACEOUS AMBER (INSECTA) 63 INTRODUCTION 63 Canadian amber perspective on the Hymenoptera 65 Comparisons to other Cretaceous amber deposits 68 GEOLOGICAL SETTING 68 Occurrence 69 Collections 69 HYMENOPTERAN DIVERSITY IN THE ASSEMBLAGE 70 Evanioidea (? Aulacidae) 70 Ceraphronoidea (Ceraphronidae, ?Megaspilidae, Stigmaphronidae) 72 Trigonalyoidea (Maimetshidae) 76 Cynipoidea (Cynipidae, Figitidae, Liopteridae, Protimaspidae) 77 Diaprioidea (Diapriidae) 79 Platygastroidea (?Platygastridae, Scelionidae) 80 Serphitoidea