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Diptera:Ephydridae AN ABSTRACT OF THE THESIS OF WAYNE N. MATHISfor the degree ofDOCTOR OF PHILOSOPHY in ENTOMOLOGY presented onApril 12, 1976 Title: A REVISION OF THE GENUS NOTIPHILAFALLEN (DIPTERA: EPHYDRIDAE) FROM AMERICA NORTH OF M..)CICO Abstract approved: Redacted for Privacy John D. Lattin The North American species of the shore fly genus Notiphila Fallen were taxonomically revised.Over 13,000 specimens were examined from throughout the Nearctic Region and information relat- ing to the biology of many species occurring in the West was gathered from both field and laboratory studies.Previously used characters were reevaluated; new evidence from comparative morphological studies on the male and female postabdomens, from developmental stages, and from biological and ecological observations was collected. Forty-nine characters were selected, quantified, and analyzed using standard numerical taxonomic procedures.Collectively, all these data were assessed and a classification derived. Forty-seven Notiphila species are now recognized from North America.These have been further classified into two subgenera and six species-groups. The subgenera were first erected by E. T. Cresson, Jr. and have been substantiated by the additional evidence accumulated during this study.Likewise, the species-groups are essentially those of Cresson with additions and minor modifications. Recognition of the subgenus Notiphila is supported by the external morphology of both sexes, by the morphology of the pupae and larvae, and by phenetic analysis using quantitative procedures. This subgenus includes 25 North American species of which three species are resurrected from synonomy (.N2 bicolor Cresson, N. cognata Cresson, and N. unicolor), two new names are proposed N. cressoni and N. paludicola for N. bicolor and N. unicolor respectively), and 13 new species are described (N. adusta, N. footei, N. pallicornis, N. taenia, N. phaeopsis, N. pulcra, N. robusta, N. eleomyia, N. latigena, N. orienta, N. pauroura, N. poliosoma, and N. shewelli).These species have been arranged into three species-groups that are based primarily on characters of the male genitalia.Most of the species of this subgenus occur in eastern North America. The second subgenus, Agrolimna, comprises 22 species, includ- ing seven new species (k_T.scoliochaeta, N. atrata, N. deonieri, N. deserta, N. elophila, N. nanosoma, and N. paludia).In addition/ the following synonymies are proposed (older name first:N. decoris Williston = N. atrisetis Cresson, N. quadrisetosa Thomson = N. occidentalis Cresson, and N. pulchrifrons Loew = N. signata Cresson). N. transversa Walker, listed as an Agrolimna species in the latest North American catalog, is a species of the genus Dichaeta. Three species-groups are recognized for the North American species, and most species are distributed in western North America. A Revision of the Genus NotiphilaFallen (Diptera: Ephydridae) from America North of Mexico by Wayne N. Mathis A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed April 1976 Commencement June 1976 APPROVED: Redacted for Privacy Profes of Entomology in charge of major Redacted for Privacy Chairan of Department of Entomology Redacted for Privacy Dean of Graduate School L.) Date thesis is presented April 12, 1976 Typed by Mary Jo Stratton forWayne N. Mathis ACKNOWLEDGEMENTS Numerous persons have assisted during all phases of thisstudy and without their contributions, much of the research could nothave been completed. Financial support was provided by National Science Foundation grant "A systematic study of the genus Notiphila Fa11n (Diptera: Ephydridae) in the Nearctic Region, " BMS 74-03135, 1974-75; by a Cana Coll Foundation grant during my visit in Ottawa (Drs.Edward C. Becker and D.M. Wood); and by a research assistantship throughthe department of entomology insect collection, Oregon StateUniversity. Type specimens were borrowed from the Academy of Natural Sciences of Philadelphia, Pennsylvania (ANSP), Drs. David C.Rentz and Selwyn S. Roback; the American Museum of Natural History,New York (AMNH), Dr. Pedro W. Wygodzinsky; the British Museum (Natural History), London (BM), Dr. Brian H. Cogan; theCalifornia Academy of Sciences, San Francisco (CAS), Dr. Paul H. Arnaud, Jr.; Cornell University, Ithaca, New York (CU), Dr. L.L. Pechuman; the Museum of Comparative Zoology, HarvardUniversity, Cambridge, Massachusetts (MCZ), Mrs. Janice C. Scott and Dr. John F. Lawrence; National Museum of Natural History,Washington, D.C. (USNM), Dr. Willis W. Wirth; Ohio State University,Columbus (OHSU), Dr. Charles A. Triplehorn; Naturhistoriska. Riksmuseet, Stockholm, Sweden (RNH), Dr. Per Inge Persson; and the Snow Entomological Museum, Lawrence, Kansas (KU), Dr. George W. Byers. The syntypes of Notiphila riparia Meigen were comparedwith Nearctic specimens through the kindness of Dr. Lo'ic Matile,Muse'um National D'Histoire Nature lle, Entomologie, Paris, France. In addition to the above, the following institutions and curators kindly loaned specimensCanadian National Collection, Ottawa (CNC), Mr. Guy E. Shewell; Colorado State University, Ft. Collins (CSU), Dr. Howard E. Evans; Florida State Collection of Arthropods, Gainsville (FSCA), Dr. Howard V. Weems; Illinois Natural History Survey, Urbana (INHS), Dr. Donald W. Webb; Iowa StateUniversity, Ames (ISU), Dr. Robert E. Lewis and Mr. Raymond Miller; Kansas State University,Manhattan (KNSU), Dr. H. Derrick Blocker; Kent State University, Kent, Ohio (KSU), Dr. B.A. Foote; Los Angeles County Museum of Natural History, California (LACM), Drs. Charles L. Hogue and Julian P. Donahue; Michigan State University, East Lansing (MSU), Dr. Roland L. Fischer; New York State Museum, Albany (NYSM), Dr. M.D. Delfinado and Mr. John A. Wilcox; Oregon State Department of Agriculture, Salem (OSDA), Mr. Richard Westcott and Mr. Kenneth Goeden; Pennsylvania State University, University Park (PSU), Dr. Ke Chung Kim; South Dakota State University, Brookings (SDSU), Dr. Edward U. Balsbaugh; University of Arizona, Tucson (UA), Dr. Floyd G. Werner; University of California, Berkeley (ICB), Dr. E.I. Schlinger; University of California, Davis (UCD), Mr. R.O. Schuster; University of California, Riverside (UCR), Mr. Saul I. Frommer; University of Georgia, Athens (UG), Dr. Warren T. Atyeo; University of Michigan, Ann Arbor (UMI), Dr. Thomas E. Moore; University of Minnesota, St. Paul (UMN), Dr. Philip J. Clausen; University of Nebraska, Lincoln (UN), Dr. Brett C. Ratcliffe; Utah State University, Logan (USU), Dr. Wilford J. Hanson; and Washington State University (WSU), Dr. William J. Turner. Plant identifications were provided by Mrs. La Rea D. Johnston, Herbarium, Oregon State University.Stereoscan electron micro- graphs were taken by Mr. Alfred Soeldner, Oregon State University. Special thanks are extended to Dr. Paul H. Arnaud, Jr. (CAS), Mrs. Janice C. Scott (MCZ), Mr. Guy E. Shewell and Dr. B.V. Peterson (CNC), and Dr. David C. Rentz (ANSP) for arranging accommodations during visits to their respective museums.I am indebted to Mr. and Mrs. George C. Steyskal, in particular, for graciously hosting me in their home while visiting the National Museum of Natural History. Mr. Steyskal also spent part of a summer in Oregon, collecting Dipterawith me and explaining the morphology of the male postabdomen.I am also appreciative of Dr. and Mrs. B.A. Foote, who arranged housing and accommodated my wife and me during a field trip to theUniversity of Montana Biological Station, Flathead Lake, Montana. Dr. CurtisW. Sabrosky, U. S. D. A. U. S. N.M. ,helped with a problem in nomenclature.His efforts are very much appreciated. I also wish to thank the members of my thesis committee, Drs. Norman H. Anderson, Kenton L. Chambers, Robert L.Dennis, Ernst J. Dornfeld, and Paul Oman.I am especially grateful and indebted to Dr. John D. Lattin, my major professor.His constant encouragement, his time, his thoughtful consideration of manyprob- lems that arose during the course of this study, and hisfriendship are sincerely appreciated. Lastly, to my wife Dianne, I am indebted for recording and organizing the locality data and for her patient encouragementthrough- out my graduate studies. TABLE OF CONTENTS Page INTRODUCTION 1 HISTORICAL REVIEW 5 METHODS AND MATERIALS 11 Materials and Generalized Procedures 11 Classification Methods and Comments 14 DISCUSSION OF TAXONOMIC CHARACTERS 17 BIOLOGY 24 CLASSIFICATION AND PHYLOGENY 33 DISTRIBUTION AND ZOOGEOGRAPHY 42 TAXONOMY 46 Genus Notiphila 46 Key to Subgenera 51 Subgenus Notiphila 52 Key to Species 55 Adusta Species-group 60 Notiphila adus to new species 60 Notiphila bella Loew 64 Notiphila biseriata Cresson 67 Notiphila cognata Cresson 70 Notiphila footei new species 73 Notiphila nudipes Cresson 75 Notiphila pallicornis new species 78 Notiphila taenia new species 81 Avia Species-group 84 Notiphila avia Loew 85 Notiphila erythrocera Loew 89 Notiphila phaeopsis new species 95 Notiphila pulcra new species 98 Notiphila robusta new species 102 Loewi Species-group 105 Notiphila carinata Loew 105 Notiphila cressoni new name 108 Page Notiphilaeleomyia new species 111 Notiphilafloridensis Cresson 114 Notiphilalatigena new species 117 Notiphilaloewi Cresson 119 Notiphilaorienta new species 124 Notiphilapaludicola new name 127 Notiphilapauroura new species 129 Notiphilapoliosoma new species 132 Notiphilashewelli new species 135 Notiphi lasolita Walker 138 Subgenus Agrolimna 143 Key to Species 147 Pulchrifrons
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