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Cretaceous Diptera from Orapa, Botswana

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Cretaceous Diptera from Orapa, Botswana CRETACEOUS DIPTERA FROM ORAPA, BOTSWANA VOLUME ONE SASKIA WATERS A DISSERTATION SUBMITTED TO THE FJI.CULTY OF SClENCE, UNIVERSITY OP THE WITWATERSRAND, JOHANNES:SURG, FOR THE DEGREE OF DOCTOR OF PHILOSOPHY. JOHANNESBURG, 1990 ii ABSTRACT An assemblage of Cretaceous, Diptera, reC!overed from the sediments of the Orapa Di",mondI1ine, Bocswana, is described. The fossil Diptera are placed In the following families: TipuJidae, Empididae, Hybotidae, Bibionidae, and possibly the Mycetophilidae, Rhagionidae and Anisopodidae. A new pulad species, Helius gotswanensis (TipuJidae, subfal(dJ.y Limoniinae), is described; it is the oldest representative of the subfamily. Ele,'en other tipuJids are described; these are not as distinctly preserved and can not be classified to lower than family level with confidence (eight probably belong to the subfamily IJimoniinae). Two new species, Empis orapaensis sp. nov. (EmpiOidae, subfamily Empidinae) and Pseudoacarterus orapaensis ge,'!. et sp. nov. (Hybotidae, slJbfarnily Hybotinae) are the earliest record of their respective subfamilies (they are the only Mesozoic members of these families from the southern hemisphere). Specimens which probably belong to the families Mycetophilidae and Rhagionidae and/or Tabanidae are described, together with the wing of a possible Rhagionid. Ten poorly preserved dipterans of 'Ilnidentified families are also recorded and illustrated; some tentative identifications include the families Mycetophilidae and Anisopodidae. Four specimens of the J:aIrdJ.y Bibionidae :florm the basis of a new genus and Ispecies, Cretobibionid;~ anticus. iii The fossils reveal details of the )alaeoenv:ironment of the Orapa region. The fa\ma lived in a crater formed by the er~ption of a Kimberlite p.ipe. 7he Diptera fed and bred near the crater lake, in a warm, humid, pxobably well- wooded or shr ubbad hab_...a.t,. with seasonal ralluall. The Oiptera were early pofklnat.ors of ang:'osperms. The concept of fossil species is discussed. 1-1any of the oipterans show remarkable stasis. iv DECLARATION I declare that this dissertation js my own ui,crlded work. It is being submitted for the degree of Doctor of Philosophy at the University of the Witwatersrand, Johannesburg. It has not been submitted before for any degree or examination in any other University. (Name of candidate) ------~----- day of -------------1990 v ACKNOWLEDGEMENTS I would like to thank the Office of the President and the National Museum of Botswana, and t:.he Debswana Mining Company for the invitation and pennission to collect fossils. The la tter also provided accommodation and financial support. Also, I acknowledge the financial SU1?Port provided by the Council of Scientific and Industrial Research and the University of the Wihtatersrand. I am grateful for the help of the following people in the collection of fcssils from Orapa: Prof. J. Kitching; Drs R.J. Rayner, J. Maguire, C. Gow, J. Masters; Mr. I.J. McKay, Mrs. M.K. Bamford, Mr. 2\. pot.ts, Mr. and Mrs. P. Forchammer, Misi5 G. Goelst, Mrs. A. Cadman, Mr. A. Cambell, Miss C. Kleinjan, Mr. P. Bender and Hr. B. Weissanbacher. My grateful and special thanks to Dr. Richard Ra.yner for endless support and supervision, and, for helpful comments and discussion, I am indebted to my friends Ian McKay and Marion Bamford. With appreciation, I wi.sh to acknowledge the help of the follo\-,'ing people for providing me with valuable information or accoass to their faciliti.es: Dr. F.S. cranston and Dr. R.I. Vane-Wright (British Museum, Natural History), Dr. W. Krzeminski (polish academy of Sciences, Krakow) and various anonymous referees and Tipulid zpeciaJJsts for providing extensive comments on the tipulid described in Chapter 2 (and Appendix 1); Dr. K.G.V. Smith (British Museum, Natural '_ vi Ristery) for checking the identification of the empid and the hybotid in Chapters 4 & 5, and, for reading and improving drafts of the manusczi.ptrs (Appendix 2 & 3) ; Dr Paul Selden (Manchester University, U.K.) and Dianne Edwards (Cardiff University), for criticism and help with the aphid (Appendix 4) and Hybotid papers (Chapter 57 Appendix 3); Drs i. Gauld and J. Cox (British Museum, Natural History) who edited the Empid paper; Dr. O.E. Heie (Institute of Biology, Royal Danish School of Educab.oniil Studies) :Eor constaructdve criticism of the aphid (Chapters & 8; Appendix 4); Dr., Judith Masters for constructive advice on Chapter 9; Dr. B. Levey (British Museum, Natural History} and D. Cullinane (Botanical Museum, Harvard University) for sending references which were unobtainable in South Africa; Dr. C. Feather of AARL for permission to use bis company's microscope; Ian McKay and Desiree Pearce and especially Dick Rayner, and, my sister Sandra Schermacher and Brian Griffin (De Beers Marine, Cape Town) j or help with the photography and loan of equdpment; and matedals; the Graphics Department at the University of the Witwatersrand for photographing most drawings. Special thanks go to my husband Keith W'aters for constructive criticism and p:r.::inting of many drafts, plotting of fig. 39, and, especially for support; and encouragement, and, Mr. A.D. Waters (Barlow Rand Computer Division, Johannesburg ) for making his photocopier available to me. vii Page Title•••.••••••••.•••.•.•••••.•••••• ,>;,..', •••••• ~ ••••••• Abstract. • •••• ••••••••••••••••••••••••••• ",. •••••••• ii Declaraticn •••••••••••••••••••••••••••••••• - ••••••••• iv AcknowIedgenents ••••••••.•••••••••••••••••.••••••••••• v Cr.)ntents of volume I: Te'xt••••••••••••••••••••••••••• vii 'Jont~~ts of volume II: Figures and Plate.s••••••• ,•••• xiii VOIllME I CHAPTER ONE - INTROlJUCl'ION 1.1. !..ocality •••••••• , ••••••••••••••••••••••••••••• 1 • 2 • Geology ••••••••••••••••••••••••••••••••••••••• 1.3. Fossil Fauna and Flora of Or,)pa••••••••••••••• 4 1.4. Methods and Materials ••••••••••••••••••••••••• 8 1.5. Dipterans as a Research Topic••••••••••••••••• 9 1.6. Introduction to the Diptera •••••••••••••• ~.••• 11 1.6.1. Diagnostic Characters of the Dipte'=a•••••• 12 1.6.2. Phylcgeny and Genealogy of the Diptera •••• 15 1.7. Introduction to the Mesozoic Insects •••••••••• ,2 1.8. Outli.~ of Contents of Chapters •••••••••••••• 22 C!1AP'I'ER TWO - SUPER!:"'AMILYTIl?ULOIDEA: TIPULIOAE 2.1. Introduction •••••••••••••••••••••••••••••••••• 25 2.2. Systematic Description and Diagnosis •••••••••• 26 viii 2. 3• Discussion....... •••••••.•••••.••••.••.••••••• 32 HeliuEJ •••............................................... .;,.......... " .... ·0 .. 'If .... 0 .. 32 Biology.•••••.••••••.••.••.•••.••••••••••••.••••. 33 Orig.ins of the Tipulidae . 34 stasis " •••••••••••••••••••• 37 SHRSand implications for evolutionary fheory •••• 39 CHAPTER THREE - O'IHER TIPt'rIDAE 3. 1. Introduction •••••••• " . 41 3.2. Descrip.tions and Discussions •••••••••••••••••• 42 3.2.1. Specimen 1••••••••••••••••••••••••••••••• 42 3.2.2. Specimen 2 . 45 3.2.3. Specimen 3 . 47 3.2.4. Specimen 4••••••••••••••••••••••••••••••• 51 3.2.5. Sr,ecimen5 . 54 3.2.6. Specim:m 6••••••••••••••••••••••••••••••• 57 3.2.7. Specimen 7••••••••••••••••••••••••••••••• 60 3.2.8. Specimen 8••••••••••••••••••••••••••••••• 62 3.2.9. Specimen 9 . 64 3.2.10. Specimen 10•••••••••••••••••••••••••••••• 67 3.2.11. Specimen 11•••••••••••••••••••••••••••••• 70 CHAPTER FOOR - SUPERFAMILY EMJ?IOOIDEA.: EMPIDIDAE 4.1. Introduction •••••••••••••.•••••••••••••••••••• 74 4.2. Systematic Description and Diagnosis •••••••••• 74 'he ix 4.3. Discussion •••••••••••••••••••••••••.•••••••.•••• 79 M tphology ••••••••••••••••••••••••••••••• , ••••••• 79 Primitive and AdV"dIlCed Features of the Venation •• 80 Biology ••••••••• < . 82 Courtship •.••••••••••••••••••••• '" •••••.••••• , •••• 84 Distribution ••••••••••••••••••••••••••••••••••••• 86 Fossil Empididae•••••••..•••••.••••••••••••••••••• B'l Evolution of hunting, feeding & courtship habits. 87 Phylogeny of the Er.~idcidea •••••••••••••••••••••• 89 5. 1. Introduction •••••••••••••••••••••••••••••••••• 91 5.2. Systematic Description and Diagnosis •••••••••• 91 5.3. Discussioo ••••••••••••••••••••••••••••••••••• 95 B~ology•••••••••••••••••••••••••••••••••••••••••• 95 Morphology••••••••••••••••••••.••••••••••••••••••• 97 Classification and Phylogeny••••••••••••••••••••• 98 Distributian •••••••••••.•••••••••••••••••••••••••• 102 CIIl\P:rER SIX - SUPERF.AMI:LY MYCEI'OPHILOIDFA: MYCEI'OI:'HILIDAE AND TlIB1\OOIDEA: IDJAGIONIDllE 6.1. Introduction •••••••••••••••••••••••••••••••••• 104 6.2. Descriptions and Dis~ussions •••••••••••••••••• 104 6.2.1. Specimen 1, description ••••••••••••.••••••• 104 Ident:!.fication ••••••••••••••••••••••••••••••• 107 Biology of the MYoatophilidae•••••••••••••••• 108 Distribution ••••••••••••••••••••••••••••••••• 110 Fossil Record •••••••••••••••••••••.••••••••••• 110 X 6.2.2. Spec:inten2" description ••••••••••••••••••• 1'11 Identification ••••••••••. , .• " •••••••••••••••• 112 Biology of the JlliagioniC!ae••••••••••••••••••• 113 Distribution •••••••••••••••••••••• .,.••••••••• 114 Fossil Record, . 114 6.2.3. Specimen 3, description ••••••••••••••••••• 116 Preservation ••••••••••••••••••••••••••••••••• 117 Identification .••••••••••••••••••••••• , ••••••• 118 Biology of the'rcbanidae ••••••••••••••••••••• 118 7. 1. In,traduction •••••••••••••••••••••••••••••••••• 120 7.2" Descriptions and Discussions . 120 7.2.1. Specinen 1••••••••••••••••••••••••••••••• 120 7.2.2. Spec:inten2••••••••••••••••••••••••••••••• 12.4- 7.2.3. Specimen 3••••••••••••••••••••••••••••••• 125 7.2.4. Spec:inten4•• eo . nJ 7.2.5. Specimen 5•••••••••••••••••••••••••••••••
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