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Proquest Dissertations INFORMATION TO USERS This manuscript has been reproduced from the microfiim master. UMI films the text directly from the original or oopy submitted. Thus, some thesis and dissertation copies are in fypewriter^ce, while others may be from any type of computer printer. The quality of this reproduction is dependent upon tfie quaiity of tfie copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6* x 9” bbck and white photographic prints are available fbr any photographs or illustrations appearing in this copy for an additional charge. Contact UMI direct^ to order. Bell & Howell Information and Learning 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0600 UMT PHYLOGENETICS AND SYSTEMAUCS OF THE WORLD STEPHANIDAE (HYMENOPTERA) Volume I DISSERTATION Presented in Partial Fulfillment o f the Requirements fbr the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Alexandre Pires Aguiar, M.S. ***** The Ohio State University 2000 Dissertation Committee: Approved by Norman F. Johnson, Adviser Wbodbridge A. Foster Hans Klompen Entomolo uate Program JohnW. Wenzel UMi Number 9971503 Copyright 2000 by Aguiar, Alexandre Pires All rights reserved. UMT UMI Microform9971503 Copyright 2000 by Bell & Howell Information and Leaming Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and Leaming Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor MI 48106-1346 Copyright by Alexandre Pires Aguiar 2000 ABSTRACT The generic relationships within the 6mily Stephanidae are examined on the basis of a cladistic analysis, the fossil species and genera are revised and its phylogeny discussed, and the Australian fauna, of importance fbr the classification of the entire fiunEy, is thoroughly investigated and revised. The relationships o f the family with other Hymenoptera is also investigated, based on a réévaluation of literature data. For the phylogenetic analysis, morphological observations are presented and discussed, and additional information and illustrations are provided for the majority of characters and their states. The analysis o f 57 species and 176 informative characters resulted in relationships and groupings which differ significantly fi'om traditional hypotheses. Results support the monophyly of the subfamily Foenatopinae Enderlein, but its original sense is expanded to mclude an aberrant new taxon, Xanthostephanus n. gen. The subâmily Stephaninae Enderlein including Schlettereriinae Orfila) is paraphyletic according to the analysis, but was maintained as a valid taxon for practical reasons. The monophyly o fthe genus Schlettererius Ashmead could not be assessed, but it is mamtained as a valid taxon based on its “distantly basal” position in relation to other stephanids. Newly discovered characters strongly support each of the following genera as u monophyletîc; Stephanus Jurine,Hemistephanus Enderlein^ Parastephanelhis Enderleiiv and Foenatopus Smith. Results suggest t h a t Benoit, Neostepfumus^<e£[tc and Diastephanus Enderieih» are not phylogenetically distinct from Foenatopus Smith, o f which th ^ are proposed as junior synonyms. The %eaasMepschus Brullé was retrieved as a distinct but highly paraphyletic lineage, composed of several independent clades. In order to preserve taxonomic stability this taxon was redefined and maintained as valid. The morphological terminology for the family is revised and illustrated, and a key and descriptions to all valid genera are provided. Analysis of combined literature data did not support the ofren-coghated sister group relationship of Stephanidae and Orussidae, suggesting instead a basal position of stephanids, together with non-ceraphronoid evaniomorphs, within the Microhymenoptera clade of a biphyletic Apocrita. The fossil o f Protostephanus ashmeadi Cockerell is confirmed to be a stephanid, and illustrated. The taxonomic status oîProtostephanus Cockerell and Electrostephanus Brues, their relationships with mctant genera of Stephanidae, and problems in defining or synonymizing fossil genera of the family are discussed. Electrostephanus is provîsorûy red^ned, two fossil species, E sulcatus sp. n. and E neovenatus sp. n. are described and illustrated from male specimens preserved in Eocene Baltic amber, and a key to species provided. The Australian stephanids are revised and k^ed. Three genera and 21 species (15 new) are recognized. The taxonomic importance of over 300 morphological characters, includmg 45 morphometric ratios, is assessed. The history of introduction and establishment o f Schlettererius cinctfpes in Australia is summarized.Stephanus in crassicauda M orl^ is transferred to Meffschusr, the female is redescribed, the male reported fer the first time and compared with the female. Occurrence of the Neotropical genus Hemistephanus in Australia is ruled out,H. giganteus Girault is synonymised with M crassicauda ^ o r l ^ ) , m A H granulatus Elliott is transferred to Parastephanellus. The groundplan o f AasXx?Siasi P arastephanellus is described in detail, and a new character set is proposed fbr taxonomic studies and descriptions of its species; P, albicepsW&oXt is proposed as a junior synonym for P. damellicus (Westwood), and P. picticeps Roman is placed as species inquirendum. New taxa areM ceneonatrix, P, albigenuis, P, adiazetos, P. brevis, P. callidictyon, P. denotatus, P, isogfyptos, P. laevigatus, P. maculatus, P. platyceratus, P. squamatus, P. strigosus, P. tanytarsus, P. trifasciatus, and P. vertipictus. Other valid species for Australia are P. caudatus Elliott, and P. ntfoom a tus (Cameron). IV ACKNOWLEDGMENTS I am deeply grateful to Norman F. Johnson, my advisor, for providing first-rate intellectual and material support to my research, and fi)r allowing me full, unconditional liberty to pursue not only my own ideas, but my own ideals, to think and materialize whatever I could, to write as much or as unconventionally as I wanted, to test, to experiment, to fail, and to succeed. For encouraging and supporting the flapping of my ambitious wings, my most smcere thank you. John W. Wenzel put the “PhJD.” in my degree. With his admirable knowledge, powerful teaching skills, and genius, John shuns away all mediocrity, allowing not only himself but also all the lucky ones around him, to aim for bright achievements under an unclouded sity. I thank him fbr so accessibily sharing his diamond talents with me. Luciana Musetti provided indispensable advice and professional help, both o f the highest quality, in nearly all parts, and fbr nearly all phases of my work: past, present, and future. Also, more sharply and intelligently tunned to the world and its people than any mommg newspaper, Luciana is an oasis of lucidity, one fiom which L fbr some luclty reason, always had the privilege o f enjoying and benefitting fi’om indefinitely. Ave, Lucianal Wbodbridge A. Foster and Hans Klompen raised the level of my committee to peak quality» and it is with pride that I acknowledge their input to my work and formation. I warmly thank Andrey Sharkov for his superb rendermg of Figure 1 (chapter 1), and for sharing with me his artistry, knowledge, witty sense of humor, and passionate eloquence. Without Jens-WOhebn Janzen the revision of the fossil stephanids would never materialize. After finding two extremely rare fossfls m this fomily, Jens kindly contacted m^ and for two years we enthusiastically cooperated in a joint paper, published while I was editing this dissertation. The holotypes of the new taxa described were made available for this study, and later donated to the %mburg museum, by Jens von Holt (Amberworld, Hamburg) (£. sulcatus sp. n.), and Carsten GrOhn (Oinde, near Hamburg) (& neovenatus sp. n.), and their kindness is also greatly acknowledged. Brian J. Armhage and Steven Passoa, although not directly involved in my work, were always receptive, always mterested, and showed such a constant and distinct aura of confidence m me, that th ^ too significantly contributed to the appearance of the final woric, and to my confidence m it. A large number of curators and other university^ scholars, listed on pages 7-13, promptly invested their time m my enterprise, and I am deeply grateful to all of them for this, hfichael J. Sharks kmdly helped with all my questions about wing venation nomenclature. Ian Naumann and Keith L. Taylor helped with several reprints of papers on Sdiiettererius cmctipeSy and Elame DeConick kindly copied and mailed me several VI important reprints of P. L. G. Benoit's papers. Theron Ellinger skillfîilly developed the films and customized all photographs that appear in this worl^ without ever frowning at any of my multiple and demanding requests. I also benefitted slgnfficantly from interacting with many other colleagues, each of which with their own admirable
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