Studies of Notiphilinae (Diptera: Ephydridae), I: Revision of the Nearctic Species of Notiphila Fallen, Excluding the caudata Group WAYNE N. MATHIS SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 287 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoo/ogy Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world cf science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 287 Studies of Notiphilinae (Diptera: Ephydridae), I: Revision of the Nearctic Species of Notiphila Fallen, Excluding the caudata Group Wayne N. Mathis SMITHSONIAN INSTITUTION PRESS City of Washington 1979 ABSTRACT Mathis, Wayne N. Studies of Notiphilinae (Diptera: Ephydridae), I: Revision of the Nearctic Species of Notiphila Fallen, Excluding the caudata Group. Smith- sonian Contributions to Zoology, number 287, 111 pages, 210 figures, 1 table, 1979. —The North American species of the shore fly genus Notiphila Fallen were revised. Over 13,000 specimens were examined from throughout the Nearctic Region, and information relating to the natural history of many species occurring in the West was gathered from both field and laboratory studies. Previously used characters were reevaluated; new characters from comparative morphological studies of the male terminalia, from development stages, and from biological and ecological observations were collected. A phylogeny is proposed based on cladistic analysis; and the phenetic relationships were assessed using standard numerical taxonomic procedures. Collectively, all these data were assessed and a classification derived. Species of Dichaeta Meigen are relegated to species-group status, but the name becomes the senior synonym of Agrolimna Cresson as one of the subgeneric names. In the subgenus Notiphila, three species-groups are recognized, comprising 25 species, of which 14 are new or required new names. In the subgenus Dichaeta, 23 species are recognized, 7 are new. Species of the subgenus Dichaeta are arrayed into four species-groups. The basis for the classification is discussed and illustra- tions of the important character states are included. The natural history and zoogeography of the nearctic species are briefly discussed. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Mantastrea cavernosa (Linnaeus). Library of Congress Cataloging in Publication Data Mathias, Wayne N. Studies of Notiphilinae (Diptera Ephydridae), I. (Smithsonian contributions to zoology ; no. 287) Bibliography: p. 1. Notiphila—Classification. 2. Insects—Classification. 3. Insects—North America. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zooloev • no 287 QL1.S54 no. 287 [QL537.E7] 591'.08s [595.774] 78-11955 ' " ' Contents Page Introduction 1 Acknowledgments 2 Materials and Generalized Procedures 3 Classification and Phylogenetic Methods 5 Discussion of Taxonomic Characters 6 Head 6 Thorax 8 Abdomen 10 Historical Review 10 Natural History 12 Tribe NOTIPHILINI Cresson 14 Key to Genera of Notiphilini 14 Genus Notiphila Fallen 14 Key to Subgenera of Notiphila Fallen 16 Subgenus Notiphila Fallen 16 Key to North American Species of the Subgenus Notiphila 17 The adusta Group 19 1. Notiphila (Notiphila) adusta, new species 19 2. Notiphila (Notiphila) bella Loew 20 3. Notiphila (Notiphila) biseriata Cresson 21 4. Notiphila (Notiphila) cognata Cresson 23 5. Notiphila (Notiphila) footei, new species 23 6. Notiphila (Notiphila) nudipes Cresson 25 7. Notiphila (Notiphila) pallicornis, new species 26 8. Notiphila (Notiphila) taenia, new species 28 The avia Group 29 9. Notiphila (Notiphila) avia Loew 29 10. Notiphila (Notiphila) erythrocera Loew 30 11. Notiphila (Notiphila) phaeopsis, new species 33 12. Notiphila (Notiphila) pulchra, new species ....'. 34 13. Notiphila (Notiphila) robusta, new species 36 The loewi Group 37 14. Notiphila (Notiphila) carinata Loew 37 15. Notiphila (Notiphila) cressoni, new name 39 16. Notiphila (Notiphila) eleomyia, new species 41 17. Notiphila (Notiphila) floridensis Cresson 41 18. Notiphila (Notiphila) latigena, new species 42 19. Notiphila (Notiphila) loewi Cresson 43 20. Notiphila (Notiphila) oriens, new species 45 21. Notiphila (Notiphila) unicolor Loew 46 22. Notiphila (Notiphila) pauroura, new species 47 iii SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Page 23. Notiphila (Notiphila) poliosoma, new species 49 24. Notiphila (Notiphila) shewelli, new species 50 25. Notiphila (Notiphila) solita Walker 51 Subgenus Dichaeta Meigen 54 Key to North American Species of the Subgenus Dichaeta, Exclusive of the caudata Group 55 The pulchrifrons Group 57 26. Notiphila (Dichaeta) pulchrifrons Loew 57 27. Notiphila (Dichaeta) hamifera Wheeler 59 28. Notiphila (Dichaeta) scoliochaeta, new species 60 The furcata Group 61 29. Notiphila (Dichaeta) bispinosa Cresson 62 30. Notiphila (Dichaeta) furcata (Coquillett) 63 The caudata Group 65 The scalaris Group 65 31. Notiphila (Dichaeta) aenigma Cresson 66 32. Notiphila (Dichaeta) atrata, new species 68 33. Notiphila (Dichaeta) atripes Cresson 69 34. Notiphila (Dichaeta) decoris Williston 70 35. Notiphila (Dichaeta) deonieri, new species 73 36. Notiphila (Dichaeta) deserta, new species 74 37. Notiphila (Dichaeta) elophila, new species 75 38. Notiphila (Dichaeta) macrochaeta Loew 76 39. Notiphila (Dichaeta) minima Cresson 79 40. Notiphila (Dichaeta) nanosoma, new species 80 41. Notiphila (Dichaeta) olivacea Cresson 81 42. Notiphila (Dichaeta) pallidipalpis Cresson 84 43. Notiphila (Dichaeta) paludia, new species 85 44. Notiphila (Dichaeta) quadrisetosa Thomson 86 45. Notiphila (Dichaeta) scalaris Loew 91 46. Notiphila (Dichaeta) sicca Cresson 93 47. Notiphila (Dichaeta) uliginosa Haliday 95 Species Inquirenda 98 Notiphila (Notiphila) latelimbata Curran 98 Notiphila (Notiphila?) riparia Meigen 98 Phylogeny and Classification 99 General Considerations 99 Tribe NOTIPHIUNI 99 Genus Notiphila 99 Subgenera Notiphila and Dichaeta 101 Species Groups 102 Subgenus Notiphila 102 Subgenus Dichaeta 104 Zoogeography 105 Literature Cited 108 Studies of Notiphilinae (Diptera: Ephydridae), I: Revision of the Nearctic Species of Notiphila Fallen, Excluding the caudata Group Wayne JV. Mathis Introduction family, especially their natural history (Brock, et al, 1969; Busacca and Foote, 1978; Collins, 1975; The family Ephydridae comprises a diverse assem- Deonier and Regensburg, 1978; Scheiring and blage of predominantly aquatic or semi-aquatic Foote, 1973; Simpson, 1976; Wirth, 1971). Despite shore flies. As larvae, most are grazers or filter- renewed efforts, however, the ephydrid fauna of feeders of organic material, especially algae and most geographic regions remains poorly known, bacteria. Others mine foliage of aquatic and terres- systematically and biologically. In the National trial plants or are saprophytic; the larvae of a few Museum of Natural History, Smithsonian Institu- species feed on spider and frog eggs. Many species tion, alone, Wirth (1968b) estimated that there are are uniquely adapted to such inhospitable environ- over 150 new species of Neotropical Ephydridae ments as sulfurous hot springs, highly alkaline or awaiting re visionary work and description. Although saline lakes, and perhaps most notably, to exposed the Nearctic ephydrid fauna is better known than pools of crude petroleum. In his book The Natural most, recent revisions of major groups (Parydra History of Flies, Oldroyd (1964:188) wrote: Stenhammar, Clausen and Cook, 1971; Hydrellia Clearly, then, Ephydridae are nothing if not versatile. Robineau-Desvoidy, Deonier, 1971; Ephydra Fallen, Evidently we are seeing in the Ephydridae a family of flies in Wirth, 1971) still required new species or new name the full flower of its evolution, and as such they offer attrac- status for approximately one-third of the taxa