NEUROPTERIDA OF THE LOWER CRETACEOUS OF SOUTHERN ENGLAND, WITH A STUDY ON FOSSIL AND EXTANT RAPHIDIOPTERA A thesis submitted to The University of Manchester for the degree of PhD in the Faculty of Engineering and Physical Sciences 2010 JAMES EDWARD JEPSON SCHOOL OF EARTH, ATMOSPHERIC AND ENVIRONMENTAL SCIENCES TABLE OF CONTENTS FIGURES.......................................................................................................................8 TABLES......................................................................................................................13 ABSTRACT.................................................................................................................14 LAY ABSTRACT.........................................................................................................15 DECLARATION...........................................................................................................16 COPYRIGHT STATEMENT...........................................................................................17 ABOUT THE AUTHOR.................................................................................................18 ACKNOWLEDGEMENTS..............................................................................................19 FRONTISPIECE............................................................................................................20 1. INTRODUCTION......................................................................................................21 1.1. The Project.......................................................................................................21 1.2. The Importance of Studying Fossil Insects...................................................22 1.3. Thesis Layout...................................................................................................23 2. LITERATURE REVIEW............................................................................................24 2.1. What are the Neuropterida?...........................................................................25 2.1.1. Neuropterida’s place within Holometabola...................................................28 2.1.2. Relationships of the three Orders...................................................................29 2.1.3. Relationships within the Orders......................................................................31 2.2. Developmental History of Neuropterida.......................................................39 2.2.1. Neuroptera......................................................................................................39 2.2.2. Raphidioptera.................................................................................................39 2.2.3. Megaloptera....................................................................................................39 2 2.3. Extant British Neuropterida...........................................................................40 2.3.1. Neuroptera......................................................................................................40 2.3.2. Raphidioptera.................................................................................................41 2.3.3. Megaloptera....................................................................................................41 2.4. Fossil Record of Neuropterida........................................................................41 2.4.1. Neuroptera......................................................................................................41 2.4.2. Raphidioptera.................................................................................................44 2.4.3. Megaloptera....................................................................................................47 2.4.4. Fossil British Neuropterida............................................................................48 2.5. Environment and Ecology..............................................................................49 2.5.1. Neuropterida...................................................................................................50 2.5.2. Neuroptera......................................................................................................51 2.5.3. Raphidioptera.................................................................................................53 2.5.4. Megaloptera....................................................................................................54 2.5.5. The Percentage of Neuropterida in Environments.........................................55 2.6. Geological Settings...........................................................................................56 2.6.1. Mesozoic flora, a brief note............................................................................56 2.7. Purbeck Limestone Group..............................................................................56 2.7.1. Palaeogeography............................................................................................56 2.7.2. Palaeoclimate.................................................................................................58 2.7.3. Palaeoenvironments.......................................................................................58 2.7.4. Lulworth Formation.......................................................................................59 2.7.5. Durlston Formation........................................................................................60 3 2.76. Palaeoflora......................................................................................................61 2.77. Palaeofauna.....................................................................................................62 2.8. Wealden Supergroup (Weald Sub-Basin).....................................................62 2.8.1. Palaeogeography............................................................................................62 2.8.2. Palaeoclimate.................................................................................................63 2.8.3. Palaeoenvironments.......................................................................................63 2.8.4. Palaeoflora.....................................................................................................65 2.8.5. Palaeofauna....................................................................................................66 2.9. Climate and Environmental Change in the Purbeck and Wealden...................................................................................................................67 2.9.1. Sedimentology.................................................................................................67 2.9.2. Palaeontological evidence..............................................................................69 3. MATERIAL AND METHODS.....................................................................................70 3.1. Fossil Acquisition.............................................................................................70 3.2. Study of the Fossils...........................................................................................70 3.3. Wing Venation.................................................................................................70 3.4. Problems Associated with Palaeoentomology...............................................72 3.4.1. Fragmentary Specimens.................................................................................72 3.4.2. The Wing Problem..........................................................................................72 PART ONE A – NEUROPTERIDA OF THE LOWER CRETACEOUS OF SOUTHERN ENGLAND: PURBECK....................................................................75 4. NEW LACEWINGS (INSECTA: NEUROPTERA) FROM THE LOWER CRETACEOUS PURBECK LIMESTONE GROUP OF SOUTHERN ENGLAND............................................76 4.1. Abstract............................................................................................................76 4 4.2. Geological Setting............................................................................................77 4.3. Material and Methods.....................................................................................78 4.4. Systematic Palaeontology................................................................................78 4.5. Discussion.......................................................................................................109 5. RAPHIDIOPTERA (INSECTA: NEUROPTERIDA) FROM THE LOWER CRETACEOUS PURBECK LIMESTONE GROUP, DORSET, UK...........................................................111 PART ONE B – NEUROPTERIDA OF THE LOWER CRETACEOUS OF SOUTHERN ENGLAND: WEALDEN.................................................................112 6. NEUROPTERAN (INSECTA) PALAEODIVERSITY WITH PREDICTIONS FOR THE CRETACEOUS FAUNA OF THE WEALDEN.................................................................113 7. NEW LACEWINGS (INSECTA: NEUROPTERA) FROM THE LOWER CRETACEOUS WEALDEN SUPERGROUP OF SOUTHERN ENGLAND..................................................114 8. TWO NEW SPECIES OF SNAKEFLY (INSECTA: RAPHIDIOPTERA) FROM THE LOWER CRETACEOUS OF ENGLAND AND SPAIN WITH A REVIEW OF OTHER FOSSIL RAPHIDIOPTERANS FROM THE JURASSIC/CRETACEOUS TRANSITION.............................................................................................................115 9. NEW SNAKEFLIES (INSECTA: RAPHIDIOPTERA) FROM THE LOWER CRETACEOUS