Bird Blow Flies (Protocalliphora) in North America (Diptera: Calliphoridae), with Notes on the Palearctic Species

Blow Flies {Protocalliphora) in North America (Diptera: Calliphoridae) j with Notes on the Palearctic Species CURTIS W. SABROSKY GORDON F. BENNETT TERRY L. WHITWORTH BIRD BLOW FLIES {Protocalliphora) IN NORTH AMERICA (DIPTERA: CALLIPHORIDAE), with notes on the Palearctic species BIRD BLOW FLIES {Protocalliphora) IN NORTH AMERICA (DIPTERA: CALLIPHORIDAE), with notes on the Palearctic species Curtis W. Sabrosky, Gordon F. Bennett, and Terry L. Whitworth Smithsonian Institution Press Washington, D.C., and London © 1989 The Smithsonian Institution Printed in the United States of America Library of Congress Cataloging-in-Publication Data Sabrosky, Curtis, W. Bird blow flies (protocalliphora) in North America (diptera: calliphoridae), with notes on the palearctic species / Curtis W. Sabrosky, Gordon F. Bermett, and Terry L. Whitworth. p. cm. BibUography: p. Includes index. ISBN 0-87474-865-8 1. Protocalliphora-North America. I. Bennett, Gordon F. n. Whitworth, Terry L. m. Title. QL537.C2423 1989 595.77 '4-dc20 89-30488 91 90 89 4 3 2 1 oo The paper used in this publication meet the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48 1984. For permission to reproduce illustrations appearing in this book, please correspond direcdy with the owners of the works. The Smithsonian Institution Press does not retain reproduction rights for these illustrations or maintain a file of addresses for photo sources. Publisher's note: For reasons of speed and economy, this book is published from camera-ready copy prepared electronically by the authors, who assume full respon- sibility for the contents and form. Dedication The authors take pleasure in dedicating this work to the many biologists, chiefly entomologists and ornithologists, who have furnished material for this study and who have been most patient in awaiting the result. In particular we honor three superb field naturalists whose interest in Protocalliphora supplied us with abundant data and material, and who encouraged and assisted in the study. We regret that Professor Spencer did not live to see the finished product. William L. Jellison, Jr. (Hamilton, Mont.) George J. Spencer, 1888-1966 (Vancouver, B.C.) Edward S. Thomas (Columbus, Ohio) Contents Introduction 1 History of the Classification of Bird Blow Flies 9 Geographic Distribution 14 The Palearctic Fauna 16 Life History 20 Do Protocalliphora Larvae Kill Young Birds? 26 Host Relations and Ecology 31 Parasites of Protocalliphora 37 Evolution and Phylogeny 37 Collecting, Rearing, and Preserving Protocalliphora 39 Control of Protocalliphora 41 Taxonomic Section Genus Protocalliphora Hough 44 Synonymy of Apaulina 46 Synonymy of Orneocalliphora 48 Status of Trypocalliphora 49 Generic Relationships of Protocalliphora 52 Taxonomy and Taxonomic Characters of the Adults 56 Identification of the Adults 65 Species Groups in the Subgenus Protocalliphora 67 Taxonomy and Taxonomic Characters of the Immature Stages 68 Key to the Nearctic Species of Protocalliphora 76 (based on males, females, and puparia) Key to Males of Nearctic Protocalliphora 81 Key to Females of Nearctic Protocalliphora 84 Key to 3rd-Instar Larvae Nearctic Protocalliphora 88 Key to Puparia Nearctic Protocalliphora 90 Descriptions of Species (alphabetical) 93 Appendix: Natural Hosts of Protocalliphora in North America 231 Annotated References 239 Index to Bird Hosts 265 Index to Protocalliphora and Miscellaneous Items 268 Figures 1-64 270 Maps 299 Preface This long-delayed study of bird blow flies {Protocalliphora) has grown steadily since its inception by the senior author about 1950 on the adult flies. Graduate study and field studies by Bennett at the University of Toronto and at the Wildlife Research Station in Algonquin Park, Ontario, provided a good foundation on the immature stages that complemented and greatly aided the study of the adults. Later graduate studies by Whitworth at Utah State University and Gold at the University of California at Albany extended knowledge of these flies for the Intermountain Region and California and added species to the known fauna. This book has been essentially ready for some years, although modified and improved from time to time, and the senior author is responsible for most of the delay. In partial extenuation, the long development of the project has undoubtedly brought us nearer to — although it has not necessarily ensured — complete knowledge of the fauna as far as the number of known species and their distribution are concerned. But any possible incompleteness does not justify further postponement, and we present our best efforts at this time as a record of the present status of knowledge and a basis for further observations. In addition to the basic taxonomic framework the book includes a review of the history and geographic distribution of Protocalliphora and a summation of other aspects such as life history, host relations, and ecology. Maps have been prepared on Goode's Base Map Series No. 102 to show the known distribution of most of the species. In this work, which deals with the species of Protocalliphora, specific names are often used by themselves to avoid too frequent repetition of the long generic name Protocalliphora or its abbreviation P. All other generic or subgeneric names are spelled in full. The host birds are referred to by their common or English names throughout the text. The Latin names are given in the list of natural hosts (Appendix), which covers all North American species mentioned as known hosts of Protocalliphora. The references contain annotations by Sabrosky on the identities of the species referred to. The book is a contribution from various organizations as follows, thus identifying the authors and acknowledging the support given them: Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, Washington, D.C. (Sabrosky). Mailing address: Systematic Entomology Laboratory, USDA, % U.S. National Museum of Natural History (NHB 168), Washington, D.C. 20560. Department of Parasitology, Ontario Research Foundation, Toronto, Ont., Canada, and (present address of author) International Reference Centre for Avian Haematozoa, Memorial University of Newfoundland, St. John's, Nfld., Canada AlB 3X9 (Bennett). A portion of the work was made possible by a research grant to the Ontario Research Foundation from the Department of Economics and Development of the Province of Ontario, and research fellowships to Bennett from the Research Council of Ontario and the National Research Council of Canada. Department of Zoology, Utah State University, Logan, Utah (Whitworth). Present address: Whitworth Pest Control Inc., 3707 - 96th St. E., Tacoma, Wash. 98446. Curtis W. Sabrosky Gordon F. Bennett Terry L. Whitworth Intnxiuction 1 Introduction Knowledge that certain calliphorid larvae ingested the blood of nestling birds dates back at least to the middle of the 19th century. Dufour (1845) in France described the hematophagous habit of the larvae in his paper on a new species that he named Lucilia dispar, nov/ considered a synonym of Protocalliphora azurea (Fallen), and Rossi (1848) reared "Musca azurea" from a large number of larvae found on young birds. In America, Walsh (1866a) mentioned large larvae found attached to the head and body of a young swallow, and he identified these as belonging to the "OEstrus family," later corrected by Osten Sacken to "Musca family in the vicinity of Musca or Sarcophaga" (Walsh 1866b). Undoubtedly these were larvae of Protocalliphora. Subsequent years saw the accumulation of many records of bloodsucking maggots on nestling birds, in both entomological and ornithological literature. Although 19th century workers made some observations, much more attention was paid to the group in the 20th century, particularly within the last few decades. The numerous past records have been discussed and summarized, in part at least, by Rodhain and Bequaert (1916), Bezzi (1922), Seguy (1929), Hall (1948), Owen (1954), Hicks (1959, 1962, 1971), and Zumpt (1965). There have been two revisions of the Nearctic species prior to the present one (Shannon and Dobroscky 1924, Hall 1948), and for the Palearctic species there have been three revisions (Zumpt 1956, Gregor and Povolny 1959, Peus 1960), a series of papers by Grunin (1966-75), chiefly on Russian species, and a revision of Trypocalliphora by Rognes (1985). Books on parasitic insects, parasitology, and birds make little or no mention of Protocalliphora. For example, there is no mention whatsoever of the genus or any of its synonyms in Marshall's recent The Ecology of Ectoparasitic Insects (1981), although there is considerable attention to such minor genera as Camus (Carnidae) and the only recently described Mystacinobia, "which may be parasitic" (Mystacinobiidae, later placed in Calliphoridae). Textbooks on ornithology and books on the biology and the life history of birds make little if any mention of Protocalliphora despite the facts that the maggots infest a sizeable proportion of birds nests, especially of passerine birds, and that they have sometimes been charged with the death of nestlings, or with weakening them so as to contribute to their death from 2 Protocalliphora other causes. In the great series by A. C. Bent (21 volumes, 1919-68) on life histories of North American birds, Protocalliphora is mentioned in only seven- -one of those only as speculation — and usually only as a brief mention or citation of one or a few references. The omission

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