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Adult and Immature Tabanidae (Diptera) of California

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Adult and Immature Tabanidae (Diptera) of California BULLETIN OF THE CALIFORNIA INSECT SURVEY Volume 22 Adult and Immature Tabanidae (Diptera) of California by WOODROW W. MIDDLEKAUFF and ROBERT S. LANE ADULT AND IMMATURE TABANIDAE (DIPTERA) OF CALIFORNIA BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 22 ADULT AND IMMATURE TABANIDAE (DIPTERA) OF CALIFORNIA WOODROW W. MIDDLEKAUFF and ROBERT S. LANE UNIVERSITY OF CALIFORNIA PRESS BERKELEY LOSANGELES LONDON BULLETIN OF THE CALIFORNIA INSECT SURVEY Advisory Editors: H. V. Daly, J. A. Powell, J. N. Belkin, R. M. Bohart, D. P. Furman, J. D. Pinto, E. I. Schlinger, R. W. Thorp VOLUME 22 Issue Date: April 1980 UNIVERSlTY OF CALlFORNIA PRESS BERKELEY AND LOS ANGELES UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND Middlekauff, Woodrow Wilson, 1913- Adult and immature Tabanidae (Diptera) of California. (Bulletin of the California Insect Survey; v. 22) Bibliography: p. lncludes index. 1. Horseflies-Classification. 2. Larvae-Insects. 3. Insects-Classification. 4. insects-California- Classification. I. Lane, Robert S., joint author. 11. Daly, Howell V. 111. Title. IV. Series: Cali- fornia Insect Survey. Bulletin; v. 22. QL475.C3C3 VOI.22 [QL537.T25] 595.7'09794s 78-66042 lSBN 0-520-09604-5 [595.7'71] lSBN 0-520-09604-5 LIBRARY OF CONGRESS CATALOG CARD NUMBER 78-66042 0 1980 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED BY OFFSET IN THE UNlTED STATES OF AMERICA Con tents Acknowledgments, vi INTRODUCTION 1 Adults, 1 Habits and Bionomics, 1 Taxonomic Treatment, 2 Key to Subfamilies and Genera of Tabanidae in California, 3 Immature Stages, 3 Habits and Bionomics, 3 Collecting and Rearing Techniques, 3 Morphology of Larvae and Pupae, 4 Taxonomic Studies, 4 Keys to Known Genera of Larval and Pupal Tabanidae in California, 6 SYSTEMATIC TREATMENT 9 Subfamily Pangoniinae, 9 Genus Apatolestes Williston, 9 Genus Brennania Philip, 14 Genus Pilimas Brennan, 16 Genus Stonemyia Brennan, 18 Subfamily Chrysopsinae, 19 Genus Silvius Meigen, 19 Genus Chrysops Meigen, 24 Subfamily Tabaninae, 39 Genus Stenotabanus Lutz, 39 Genus Haematopota Meigen, 40 Genus Atyfotus Osten Sacken, 41 Genus Hybomitra Enderlein, 42 Genus Tabanus Linnaeus, SO Literature Cited 59 Plates 1-15 65 Index to Tabanid Species and Synonyms 97 Acknowledgments Acknowledgment is made to the following individuals J. R. Anderson, University of California, Berkeley, for and institutions whose assistance made possible the helping with the 1973 parity determinations of Chrysops study of institutional collections in their care. The ab- hirsuticallus and for many other kindnesses during the breviations below, and in the text where collection data course of this study; V. F. Lee, California Academy of are given, indicate the institution where specimens Sciences, San Francisco, for providing us with larvae of studied are to be found: C. B. Philip, P. H. Arnaud, Apatolestes actites and for participating in a few of the Jr., California Academy of Sciences, San Francisco field trips; A. H. Murphy, Superintendent, University (CAS); C.L. Hogue, Los Angeles County Museum of California, Hopland Field Station, for use of the (LACM); S. I. Frommer, University of California, facilities; J. L. Sansing, Superintendent, Point Reyes Riverside (UCR); W. M. Rogoff, United States Depart- National Seashore, for permission to study the tabanid ment of Agriculture, Fresno (USDAF); M. s. Was- fauna of the Seashore; Pamela Vesterby, Lauren G. bauer, Bureau of Entomology, California Department Zarate, and Celeste Green for their drawings of adults of Agriculture, Sacramento (CDA); E. I. Schlinger, and immatures; J. Fraser, Department of Entomologi- J. R. Anderson, J. A. Powell, California Insect Survey, cal Sciences, University of California, Berkeley, for Department of Entomological Sciences, University of photographs of the wing patterns; Susan J. Lane for California, Berkeley (CIS); R. 0. Schuster, University determining in part the hatching percentages and para- of California, Davis (UCD); L. L. Pechuman, Cornel1 sitization rates of eggs; and G. Grodhaus, Vector and University, Ithaca (CU); United States National Mu- Waste Management Section, California Department of seum (USNM); California Department of Health, Health, for taxonomic advice and editorial suggestions. Berkeley (CDHB). We gratefully acknowledge assistance from individ- We thank H. J. Teskey, Biosystematics Research In- uals in the Department of Entomological Sciences, stitute, Agriculture Canada, for his helpful comments University of California, Berkeley, who identified vari- on the sections on biologies and immature stages and ous taxa of arthropods and plants, including M. Bent- for sending us larvae of Haematopota americana; J. F. zien (spiders), L. E. Caltagirone (Scelionidae), J. T. Burger, Department of Entomology, University of New Doyen (invertebrates and plants associated with pango- Hampshire, for reviewing part of the manuscript; C. B. niine larvae), K. S. Hagen (Coccinellidae), and P. Philip for companionship during several of the field Rubtzoff (plants). investigations and numerous ideas and suggestions; INTRODUCTION1 Horse flies and deer flies are economically important involved. The notes emphasize features which will aid insects. Adult females of many species suck blood; their the reader in species identification. approach annoys potential victims and they may inflict Information presented on adult taxonomy and distri- a painful bite. Some species transmit mechanically, or bution was prepared by Middlekauff; the biosystematic in a few cases biologically, pathogenic agents such as studies of the immature stages and the biology sections arboviruses, bacteria, protozoa, and nematodes. Fe- were prepared by Lane, unless otherwise noted. males of some Central and South American species carry eggs of the human bot fly, Dermatobia hominis (Linnaeus, Jr.), which causes cutaneous myiasis on a ADULTS suitable avian or mammalian host. The bacterial dis- eases, tularemia and anthrax, are also transmitted by Adult North American tabanids vary in size from the bite of tabanids. For a comprehensive review of the about 6.5 to 30 mm in length. These usually stout, big animal disease agents transmitted by tabanids see headed, and big-eyed flies are found from sea level to Krinsky (1976). 3,300 m or more. Since the first treatment of California Tabanidae by Middlekauff (1950) some taxonomic problems have Habits and Bionomics been resolved; 9 new species have been described and Tabanids frequent a great variety of habitats, rang- there have been several new state records; new data on ing from deserts to alpine environments, usually with a geographical distribution and seasonal patterns have moist or wet breeding site nearby. Most are active on been acquired; and there has been an impressive in- warm, sunny days. Flight activity decreases when it is crease in our knowledge of immatures and the habits of cool, breezy, or both. There are notable exceptions adults. along the California coast where Brennania heru (Osten In California there are 73 species and 7 subspecies of Sacken) and Apatolestes actites Philip and Steffan are Tabanidae in 11 genera. The genera Chrysops, Hybo- active under both cool and windy conditions. Some spe- mitra, Apatolestes, and Tabanus contain the most spe- cies are rare, whereas others may occur in such num- cies with 21, 13, ll, and 10 respectively, which account bers that their persistent and painful attacks make nor- for 75 percent of all species recorded from the state. mal outdoor activities for humans, domestic livestock To save space, we have eliminated detailed listing of and mammalian wildlife unbearable or uncomfortable. collection data except for the less common species. For Feeding habits of Tabanidae vary. Most females are the more common species, we either list the counties or haematophagous, but some. as well as all males, subsist give a summary of the distribution pattern. The ap- on nectar and plant exudates. No tabanid is known to proximate collection localities of all specimens studied be host specific, although definite preferences are are shown on topographic maps. Seasonal flight periods shown by some species. Most are diurnal feeders, al- are also summarized for each species. All references to though some feed at dawn, dusk, or at night, particu- time pertain to local time as far as we are aware. larly in the tropics. The descriptive notes in the discussion section are not The haematophagous members of the family are intended to be a taxonomic redescription of the species known to favor larger mammals as hosts. Members of the horse (Equidae), cow (Bovidae), deer (Cervidae), 1. Supported in part by Experimental Station Project No. 3502. and camel (Camelidae) families seem to be preferred 1 2 BULLETIN OF THE CALIFORNIA INSECT SURVEY hosts, but many other mammals as well as birds, liz- taken from vegetation or the ground near the larval ards, and turtles also are attacked. On rare occasions, habitat or mating site. Males, with rare exceptions, are tabanids have even been observed feeding on dead collected infrequently. Best results are attained by mammals. Many workers have noted that the Tabani- sweeping flowers, grass, or other vegetation near emer- nae generally prefer larger hosts than do the Chrysop- gence sites, on sun-lit wooden walls and fences, or when sinae (Philip, 1931; Stone, 1930; MacCreary, 1940; they are hovering or resting near mating sites. Forest Pechuman et ai., 1961; Smith et ai., 1970), and that clearings, pathways, country roads, or hilltops are all humans are more often troubled
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