The Morphology and Taxonomic Value of Thoracic Structures in Some Brachycera, Diptera
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\ THE MORPHOLOGY AND TAXONOMIC VALUE OF THORACIC STRUCTURES IN SOME BRACHYCERA, DIPTERA by MUSA ABDALLA AHMED, D.I.C., M.Sc. (London) Thesis submitted for the degree of Doctor of Philosophy of the University of London Department of Pure and Applied Biology, Imperial College of Science and Technology, South Kensington, S.W.7. July 1982 jXJrl JjLJ' J& -^llUT J^ ^ l^r tLe^Vf f Jfc'iej _xx»£x x . - -- x x x» xxx x » > • > x x * i — x> x LiJcU ^LJ Ij|U Cn) ^O^JlA i- - >lxfl —£xx » —X»t f X x x XX > /» . > x»r x I x S ^UIUA ^Ur-u ^^^J^^lib JU eg) ^-^IJ^T^UJT vil;^ x x^xvix ».x xx £ „ X »x >x»l v £ »xl xx » j^ju-U^lj iU JiU' JU ^tH- X > XX (g) O^xj^TUj rr'-n . iyM1 <T> /r? f/ie name o/ God, f/?e Merciful, the Mercy-Giving He taught Adam all the names of everything; then presented them to the angels, and said: "Tell me the names of these, if you are truthful." They said: "Glory be to You; we have no knowledge except what You have taught us. You are the Aware, the Wise!" He said: "Adam, tell them their names." Once he had told them their names, He said: "Did I not tell you that I know the Unseen in Heaven and Earth? I know whatever you disclose and whatever you have been hiding." The Cow 2: 31-33 THE MORPHOLOGY AND TAXONOMIC VALUE OF THORACIC STRUCTURES IN SOME BRACHYCERA, DIPTERA ABSTRACT The thoracic morphology of some Brachycera (Diptera) is considered. The results are presented in descriptions of the thoracic skeletons of 19 species from 19 families, to- gether with 76 text figures, and a systematic survey of major families. Homologies of parts are suggested and a uniform morph- ological terminology is used for these. The possible use of some exoskeletal and endoskeletal structures in Brachycera systematics is assessed. Among those features given particular attention are the sella (presternum), posterior intersegmental suture and probasisternum (prosternum) . The Asilids possess a sella which is sufficiently distinct both to unite them as a monophyletic group and to separate them from other Brachycera. The shape of the sella readily divides the Bombyliidae into two major groups, Anthracinae and Bombyliinae sensu lato, thus confirming the broad trad- itional classification of this family. The state of the posterior intersegmental suture can be reliably used for recognition of the Empidoidea as a discrete and natural group. The condition of probasisternum, whether or not united to the proepisternum, helps to define subfamily limits in Empididae, Some genera have been found to possess a prephragma of a characteristic shape, but these shapes may occur independ- ently in different groups. Of the peculiar features demonstrated by Systropus edwardsi (Bombyliidae) the following are reported for the first time: the profurcal pits are confluent, the prospina- sternum is present and gives rise to a well developed spina internally, and the metafurca extends in the thoracic cavity as a single, cup-shaped structure. I - iv - ACKNOWLEDGEMENTS This study has been conducted under the supervision of Mr. H.E. Goto, to whom I am most grateful for many useful suggestions and. for critically reading the manuscript. I have been fortunate during the course of my work to benefit from the extensive knowledge and experience of Professor R.G. Davies, especially in matters concerning nomenclature and homologies. Donation of specimens from the collections of the British Museum (N.H.) has greatly facilitated my work. For this I am indebted to members of staff of the Diptera Section. Special thanks are due to Mr. B. H. Cogan for his never-fading consideration and for selflessly attending to every request for specimens. I owe a debt of gratitude to Prof essor J. Bowden of Rotham- stead Experimental Station for providing material from his own collection, for his stimulating conversations and for his interest in the project. The most generous gifts of specimens came from the private collections of Dr. J. Wilcox of Anaheim, California and Mr. C. E. Dyte of Slough Laboratory, to both of whom I am extremely grateful. I also appreciate the help of those who have sent me specimens and/or literature. They are: Dr. H. Ulrich, Museum Alexander Koenig, Bonn; Dr. D. W. Webb, Illinois Institute of Natural Resources, Champaign; Dr. W. W. Worth, U.S. National Museum, Washington; Professor L. L. Pechuman, Department of Entomology, Cornell University, Ithaca; Dr. R. Matsuda and Dr. H. J. Teskey, Biosystematics Institute, Ottawa; Dr. J. Waage, Imperial College Field Station; Dr. J. G. Londt, Natal Museum; Mr. M. A. Cotton, R.C.S. London,; and the late Osman H. Adlan, Agricultural Research - V - Corporation, Wad Medani, Sudan. I acknowledge Elizabeth Jenks for her patience in typing the thesis. Thanks to the authorities of the Agricultural Research Corporation, Wad Medani, Sudan, for financial support. - vi LIST OF FIGURES FiPage 1. Lateral view of the thorax of Atylotus agrestis with details of sutures 78 2. Lateral view of the thorax of A. agrestis with details of sclerites 79 3. Lateral internal view of the thorax of A. agrestis 80 4. Sternum of A. agrestis 81 a. ventral view b. dorsal internal view 5. Ventral view of the probasisternum in Tabanidae 82 a. Haematopota patellicornis b. Chrysops brucei 6. a. Posterior internal view of the prephragma of 83 A. agrestis b. Posterior internal view of the metafurca of A. agrestis 7. Lateral view of the thorax of Pelecorhynchus fusconiger 8. Lateral view of the thorax of Pantophthalmus tabaninus 85 9. a. Ventral view of the probasisternum of 86 Pelecorhynchus fusconiger b. Ventral view of the probasisternum of Pantophthalmus tabaninus 10. Posterior internal view of the prephragma 87 a. Pelecorhynchus fusconiger b. Pantophthalmus tabaninus 11. Lateral view of the thorax of Chorisops tibialis 88 12. Lateral internal view of the thorax of C. tibialis 89 - vii - Fig. Page 13. Sternum of C. tibialis 90 a. ventral view b. dorsal internal view 14. Ventral view of the probasisternum in Stratiomyidae 91 a. Beris chalybeata b. Merosargus stamineus 15. Ventral view of the probasisternum in Stratiomyidae 92 a. Stratiomys chamaeleon b. Hermetia illucens 16. Posterior internal view of the prephragma in Stratiomyidae 93 a. Chorisops tibialis b. Pachygaster atra 17. Posterior internal view of the prephragma in Stratiomyidae 9A a. Hermetia illucens b. Merosargus stamineus 18. a. Lateral view of the thorax of Xylomya tenthred- inoides 95 b. Lateral view of the thorax of Xylophagus fulgidus 19. Ventral view of the probasisternum 96 a. Xylophagus fulgidus b. Xylomya tenthredinoides 20. Posterior internal view of the prephragma 97 a. Xylophagus fulgidus b. Xylomya tenthredinoides 21. Lateral view of the thorax of Coenomyia ferruginea 98 a. external b. internal 22. a. Posterior internal view of the prephragma of ' Fekruginea 99 b. Ventral view of the probasisternum of C. ferruginea - viii - Fig. Page 23. Lateral view of the thorax of Rhagio scolopaceus 100 24. Lateral internal view of the thorax of R. scolopaceus 101 25. a. Lateral view of the thorax of Hilarimorpha sidora 102 b. Lateral view of the thorax of Vermileo sp. 26. Probasisternal variation in Rhagionidae 103 a. Rhagio scolopaceus b. Atherix ibis c. Chrysopilus cristatus d. Vermileo sp. 27. Prephragmal variation in Rhagionidae 104 a. Rhagio. scolopaceus b._ Atherix ibis c. Chrysopilus cristatus d.. Vermileo sp. 28. a. Posterior internal view of the prephragma of Hilarimorpha sidora 105 b. Ventral view of the probasisternum of H. sidora 29. Lateral view of the thorax of Exhyalanthrax lugens 106 30. Lateral internal view of the thorax of E. lugens 107 31. Thorax of Systropus edwardsi 108 a. Lateral view b. Posterior view of the metathorax 32. Ventral view of the cervix and probasisternum 109 a. Exhyalanthrax lugens b. Bombylius major 33. Ventral view of the cervix and probasisternum a. Enica longirostris b. Callostoma fascipennis - IX - Fig. Page 34. Probasisternal variation in Bombyliidae 111 a. Heterotropus indicus b. Systropus edwardsi 35. Prephragmal variation in Bombyliidae 112 a. Bombylius major b. Bombylius discoideus c. Callostoma fascipennis d. Usia aenea e. Toxophora maculata f. Phthiria gaedii 36. Prephragmal variation in Bombyliidae 113 a. Exhyalanthrax lugens b. Exoprosopa minos c. _ Villa sexfasciata d. Heteralonia megerlei e. Tomomyza pictipennis f. Anthrax pithecius 37. Pterothoracic furca in Bombyliidae 114 a. Systropus edwardsi b. Exhyalanthrax lugens 38. Metafurcal variation in Bombyliidae 115 a. Bombylius discoideus b. Bombylius major c. Exoprosopa aurulans d. Exoprosopa louisae 39. Metafurcal variation in Bombyliidae 22.6 a. Usia aenea b. Toxophora maculata c. Phthiria gaedii d. Tomomyza pictipennis 40. Lateral view of the thorax of Ogcodes gibbosus 41. Ventral view of the thorax of Qgcodes gibbosus hq - IX - Fig. Page 42. Probasisternal variation in Acroceridae 119 a. Eulonchus tristis b. Acrocera globulus 43. a,b-prephragmal variation in Acroceridae 120 a. Ogcodes gibbosus b. Acrocera globulus c- Posterior internal view of the metafurca of Ogcodes gibbosus 44. Lateral view of the thorax of Prosoeca accincta 121 45. a. Posterior internal view of the prephragma of Prosoeca accincta 122 b. Ventral view of the probasisternum of Prosoeca accincta 46. a,b- Probasisternal variation in Nemestrinidae 123 a. Fallenia fasciata b. Neorhynchocephalus sulphureus Ch Posterior view of the postphragma of Prosoeca accincta 47. Lateral view of the thorax of Ectyphus pinguis 124 48. Lateral view of the metathorax of Mydas clavatus 125 49. a,b- Prephragmal variation in Mydidae 126 a. Ectyphus pinguis b. Mydas clavatus c- Ventral view of the probasisternum of Ectyphus pinguis 50. Lateral view of the thorax of Scenopinus fenestralis 127 a. External b. Internal 51. a. Posterior internal view of the prephragma of ]_2g Scenopinus fenestralis b.