Bird Blow Flies (Protocalliphora)

in North America (Diptera: Calliphoridae)

. with Notes on the Palearctic Species

CURTIS W. SABROSKY | GORDON F. BENNETT TERRY L. WHITWORTH

BIRD BLOW FLIES (Protocalliphora) INNORTH 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: cal- liphoridae), with notes on the palearctic species / Curtis W. Sabrosky, Gordon F. Bennett, and Terry L. Whitworth.

p- cm.

Bibliography: p.

Includes index.

ISBN 0-87474-865-8

1. Protocalliphora--North America. I. Bennett, Gordon F. Il. Whitworth, Terry L. III. Title.

QL537.C2423 1989

595.77°4--de20 89-30488

Ot 907.89 A SK 2a

co 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 directly 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

MANEHOMUCTIOME taievad ech wie eh ira, ears lem adrian tas amie sab arta Ao, ] History of the Classification of Bird Blow Flies ........ 9 GcosraphniceDistributiony 25) os es 14 ihe mealearctic FauUmMay. ci es cu ala! cei aa uaan sed omaiis eelane 16 IPC METIS CORY err etre tse ee Ut ets Beal UR Sits odie nee a 20 Do Protocalliphora Larvae Kill Young Birds?.......... 26 Mostuelationstand Ecology 75 4. ea eons 31 ALASKtESHOLMEFOLOGCQHITDNONG 2.00 oh ee eon te eee 37 Evolutions and Phylogemyiis oo ee in es oe ee 37 Collecting, Rearing, and Preserving Protocalliphora ..... 39 Control Of ProrocglhlipRora. oO ee te Cn oe oe 4] Taxonomic Section

Genus-Protocalliphora Hough’ 8 eo. en ee 44 SVMOMN MY Of AD QITO I aie r ie een ities Se aia 46 Synonymy olf Orneocalliphora ie! oo ee es 48 SAtUSHOL MP POCAIITDHONG: ase face ent enon snes. 49 Generic Relationships of Protocalliphora ............. 52 Taxonomy and Taxonomic Characters of the Adults ..... 56 dentification.of the Adults) 2002). .35) Rk ee 65 Species Groups in the Subgenus Protocalliphora ........ 67 Taxonomy and Taxonomic Characters of the

TTT EUTE STAB SS ek a an a ae es Nance eIMIn Aarne Screnarts 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

INOrthvAMeriCa es! rie ee son ee ene er ple Meas ie eNe ue ot 231 Annotated References ane, gait se ona 239 Indexto Bird Hosts 3c OE Ge oe) 265 Index to Protocalliphora and Miscellaneous Items ...... 268 PSUs OA ee he ey ORM Ay aa OME ime ey ae vias ee 270

LOSE perk ap ne Ne OAR ot REE OIC Zee 0 a ae AO RCN TRUM MRR RC DR 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 A1B 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

Introduction |

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, now considered a synonym of Protocalliphora azurea (Fallén), 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), Séguy (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 Carnus (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 or scanty mention by ornithologists seems especially curious in view of the numerous papers by C. W. Johnson in ornithological journals, the extensive rearings reported by Shannon and Dobroscky (1924), and the papers of Henshaw, Mason, McAtee, C. W. Miller, Plath, Stoner, Storer, and others, which antedated most of Bent’s volumes and the recent books and texts on birds and their lives. In the periodical literature of ornithology there is rarely more than a mere mention of the bird blow flies and their maggots; Protocalliphora, if mentioned, is commonly not even indexed. Even papers specifically on the insect faunas of birds nests all too often have lists of lice, fleas, mites, and ticks with no mention of Protocalliphora. Wing’s (1956) book on the Natural History of Birds: A Guide to Ornithology is one of the few bird books to say much about Protocalliphora, but even so this is not indexed.

A common name is perhaps unnecessary for these flies. The adults are seldom net collected, and the maggots are usually found by entomologists, for whom the generic name Protocalliphora is readily understood, or by ornithologists. Various publications have used such names as "blood-sucking — larva flies," "parasitic bird-flies," "nestling screwworms," and "bird-nest screwworms," but these seem awkward. The last two are also misleading because the larvae do not have the characteristic appearance of screwworms. For that matter, "blow fly" is also inaccurate because Protocalliphora larvae do not "blow" carcasses, but it is the common name for flies of the family Calliphoridae. The German literature uses the appropriately suggestive name "Vogelblutfliegen" (bird blood flies), although this might imply, incorrectly, that the flies themselves suck blood, and the term would also fit some Hippoboscidae. There are also other flies with ornithophilic blood-sucking larvae, such as Philornis (Muscidae) and Neottiophilum (Neottiophilidae). But if a common name is desired, "bird blow flies" would be short and euphonious; blow flies (calliphorids) that attack birds. This was adopted by Zumpt (1965).

Difficulties in the identification of Nearctic Protocalliphora stimulated Sabrosky about 1950 to reexamine the classification of the genus. Soon after, Bennett, based on observations by D. M.

Introduction 3

Davies in 1949, undertook a study of the ecology, life history, and immature stages of the genus, principally at Lake Sasajewan (45° 35’ N., 78° 30’ W.) in southern Algonquin Park, Ontario, at the Wildlife Research Station of the Ontario Department of Lands and Forests. The parallel studies on adults and immature stages, although initially not equally comprehensive, proved to be so mutually advantageous and complementary that they soon developed concomitantly. The ecological and life history studies have also contributed significantly to understanding of the genus. These are summarized from an unpublished doctoral dissertation (Bennett 1957) that was generously made available to other workers and has been cited by them, and from the MS. thesis and doctoral dissertation by Whitworth in Utah (Whitworth 1971, 1976, 1977).

As a result of our studies, covering the extensive Algonquin Park and Utah material, plus collections from all major museums in this continent and other sources in the United States and Canada, the present revision recognizes 26 species (15 new and one newly recognized as Holarctic), compared with 10 recognized by Hall (1948) in the last revision of the genus in North America. Furthermore, unlike most previous publications, immature stages are also covered. Table I presents a summary that shows the stages that are known and described in this paper. The studies are based on a total of 9,292 adult specimens (4,649 males, 4,643 females), and large numbers of larvae and puparia. Of the 16 names already published and available, the senior author has studied the types of 13, one is lost, and the other two are European names now being used in North America.

Much remains to be learned about Protocalliphora, in spite of the nearly complete record for mature larvae, puparia, and adults (Table I). The biology and ecology of most species are unknown, at most inferred from host records. The complete life history has not been observed for any species. The distribution of most of the species is spottily known, although the scattered records indicate wide ranges. Presumably, Protocalliphora occurs throughout the nesting ranges of the bird hosts, with the exception of the far south as judged from the typically northern or higher altitude range of the genus (cf. Map 1). However, a few specimens from Mexico and southwestern United States suggest that attention to the Sonoran fauna would yield interesting results. Few areas have been studied thoroughly, and isolated available specimens hint at other undescribed species.

4 Protocalliphora

It is apparent from some puzzling complexes that much work remains to be done before the far western fauna is fully understood. Certain species, such as asiovora and chrysorrhoea,

Table I. The described stages of Nearctic Protocalliphora

Species Egg Larval Instars Puparium Adults Ist 2nd 3rd Male Female

Subgenus Trypocalliphora braueri

Subgenus Protocalliphora aenea xX asiovora avium »,« >< beameri bicolor brunneisquama chrysorrhoea cuprina deceptor fallisi halli hesperia hesperioides hirundo xX interrupta lata metallica xX Xo parorum sapphira seminuda shannoni sialia x spatulata spenceri =x tundrae =X

KKM He eM eM MM OM * * *

+ ~ KKK KKK lel i elelelelalelalalalvia xxKmK

x ~~

xX Xx X

KK PK KP KKM MMMM OM KP PP KKK OM

* = Important characters described from puparium. ! = Atypical; only two undersized puparia known.

Introduction 5

are clearly marked. In species such as hesperia, however, one finds it difficult to determine whether one is dealing with one or a few variable species in which the local populations differ slightly, or whether there are a number of closely related species. Such factors as mixed infestations, lack of outstanding characters in both sexes, lack of conspicuous differentiation in the immature stages, and immaturity of many reared series make the interpretation extremely difficult.

Perhaps some hybridization is taking place, or the western region is one of active speciation and the local populations show tendencies that have not yet stabilized as_ consistent characteristics. On the other hand, reasonable stability may be present, but obscured by unfortunately narrow distinctions between species that may occur together in mixed infestations. Purposeful field work like that of Bennett in the Algonquin Park area, Whitworth in Utah, and Gold and Dahlsten in California is essential to clarify the distinctions and relationship. In the meantime, a tentative arrangement is offered as the best that can be deduced at this time from the available material. Certainly in the West it is even more necessary than in the East that firm identifications be based on series containing both sexes, plus mature larvae or puparia whenever possible. Separation of these western forms from the numerous known species will probably be difficult and will require increasingly careful attention to details.

Keys and detailed descriptions are given for males, females, and immatures. For each species there is a brief statement of the known distribution, followed by the detailed records. For already described species, the identification records have been somewhat abbreviated, if numerous; the detailed records will be filed with the Systematic Entomology Laboratory, USS. Department of Agriculture, Washington, D.C., or in the Archives of the Smithsonian Institution, so that future workers may locate and recheck the material if later developments require it.

Voucher specimens for published records have been checked whenever possible, and unconfirmed published records are few in number. For a few species, such as the occurrence of avium in crows’ nests and in the ears of hawks, or records of braueri (hirudo) reared from larvae found feeding subcutaneously, the records can be accepted as almost certainly correct.

Dates of reared specimens are of little significance for Protocalliphora. Life history studies show that the breeding of Protocalliphora species is confined to and continuous throughout the nesting seasons of their bird hosts. Moreover, dates on most

6 — Protocalliphora

reared specimens do not state whether they are the dates of nest collection or of emergence of adults. Where dates are given with host records, they are those of emergence of adult flies unless otherwise specified. Dates of hand-caught specimens may be significant, however; early and late dates give evidence of overwintering by adults.

The collector’s name, if known, is stated for type series and for uncommon species, but not for the numerous records of common species already described, such as aenea, avium, and sialia. The collector’s name is sometimes supplied from published information, notably from the papers of C. W. Johnson (1925-32).

For each species there is a list of known hosts and a brief statement of the known or inferred ecology. The latter is based on the field studies plus analysis of other available records and of published information where the species identifications have been checked. Ecological data on the host birds have been derived from the publications by Bent (1919-53), Peterson (1947), and Taverner (1934), as well as the field studies by Bennett and Whitworth.

Host records refer to flies reared from larvae and puparia found in the nests of the hosts, but any records of direct attack upon the birds themselves, e.g., myiasis in the ears, or larvae in abscesses under the skin, are so specified and given in the detail available.

Location of paratypes and other material is indicated by the following abbreviations for the frequently cited collections:

CAS California Academy of Sciences, Golden Gate Park, San Francisco, Calif. 94118

CNC Canadian National Collection, Agriculture Canada, Ottawa, Ontario. KIA OC6

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 01238

UBC University of British Columbia, Spencer Entomological Museum, Vancouver, B.C. V6T 2A9

USNM_ USS. National Museum of Natural History, Washington, D.C. 20560

USU Utah State University, Department of Biology, Logan, Utah. 84322

Infrequently cited collections have either been cited in full or somewhat shortened by standard and readily understood

Introduction 7

abbreviations, e.g., U. Del. = University of Delaware; U. Calif., Berkeley = University of California at Berkeley. University material is usually in the collection of the Department of Entomology.

Acknowledgments

The writers gratefully acknowledge their indebtedness to numerous individuals and institutions for the loan of material and for other help during the course of this study. Particular thanks are due to A. Murray Fallis, former Director, Department of Parasitology, Ontario Research Foundation, Toronto, Ont., and Wilford J. Hanson, Department of Biology, Utah State University, Logan, Utah, under whom Bennett and Whitworth, respectively, carried out field and laboratory investigations for their doctorate programs, for their sympathetic interest and sound counsel; to the Ontario Department of Lands and Forests for making available the facilities of their Wildlife Research Station, Algonquin Park, Ontario, and to the staff and other colleagues at the Station for assistance and information during the field work; to D. M. Davies, McMaster University, for initiating the field studies in Algonquin Park; to Kenneth J. Capelle, Brigham City, Utah, (retired, Fish and Wildlife Service, U.S. Department of the Interior), for suggesting Whitworth’s study and for his assistance throughout the project; to the late G. J. Spencer, University of British Columbia, for the generous and long-time loan of his large collection of Protocalliphora gathered over a period of some years; to W. L. Jellison, Rocky Mountain Laboratory, U.S. Public Health Service, for making available the large collection of Protocalliphora accumulated by himself, C. B. Philip, and others in the course of their field work; to the late J. C. Bequaert, Museum of Comparative Zoology, Harvard University, and Arthur G. Humes, Boston University, for the loan of important collections accumulated and published on by Charles W. Johnson; to Edward S. Thomas, Ohio State Museum, the late M. T. James, Washington State University, E. L. Kessel and the California Academy of Sciences, and Clifford S. Gold and Donald L. Dahlsten of the University of California at Albany, for the loan of particularly important collections; to George E. Watson, Smithsonian Institution, and Richard Banks, Fish and Wildlife Service, U.S. Department of the Interior, for many courtesies in connection with ornithological literature and the nomenclature of the bird hosts, and to Knut Rognes of Norway for generously sharing with us his results of dissections

8 Protocalliphora

of types and his descriptions from his manuscript on Calliphoridae for the Fauna Entomologica Scandinavica.

We are indebted to Deborah Roney for the drawings and the makeup of the plates, to Richard Banks, Michael Schauff, Guy E. Shewell, and Thomas Pape for careful and critical reading of the manuscript and excellent comments, to two anonymous referees, to Vera Lee for typing of the manuscript, to Elizabeth Klafter for the preparation of camera ready copy, and to Linda Lawrence for adjusting plates to final copy.

Our appreciation for material and information is also due to the following individuals, some of whom are deceased: John F. Anderson, Paul H. Arnaud, Jr., John R. Baker, Russell P. Balda, Sister Barbara Ann (All Saints’ Convent, Catonsville, Md.), R. H. Beamer, W. W. Becklund, John N. Belkin, Robert Bohm, Gary R. Bortolotti, J. W. Boyes, A. E. Brower, George W. Byers, Robert A. Cannings, S. G. Cannings, K. J. Capelle, John A. Chapman, Frank R. Cole, B. E. Cooper, Monique Coulloudon, Scott Crocoll, C. H. Curran, Vivie E. Davis, Vasiliki Demas, Henry Dietrich, H. R. Dodge, Richard P. Dow, W. L. Downes, Jr., Emmet R. Easton, Howard E. Evans, W. G. Evans, D. C. Ferguson, R. H. Foote, Woodbridge A. Foster, Norman R. French, John George, J. D. Gregson, G. C. D. Griffiths, A. H. Grewe, Jr., K. J. Grunin, Harvey L. Gunderson, G. E. Haas, David G. Hall, Jr., Jeffrey A. Halstead, Mrs. Frances Hamerstrom, Joseph J. Hickey, Cluff Hopla, Donald S. Horning, Jr., Paul D. Hurd, Jr., C. D. Johnson, W. W. Judd, Rokuro Kano, Ben Keh, E. E. Kenaga, Ke Chung Kim, Frank Kuhlman, W. E. LaBerge, Paul K. Lago, Robert D. Lee, Hugh B. Leech, Robin Leech, B. Lindeberg, Tom Lund, J. E. H. Martin, Wayne N. Mathis, J. F. McAlpine, A. T. McClay, E. T. McKnight, John F. Mehner, Heinz Meng, Donald H. Messersmith, Virgil I. Miles, Robert T. Mitchell, Val Nolan, Jr., Pekka Nuorteva, Don R. Oliver, L. L. Pechuman, P. I. Persson, Fritz Peus, T. David Pitts, Adrian C. Pont, Kim G. Poole, Dalibor Povolny, H. W. Prescott, Frank W. Preston, Robert L. Rausch, H. J. Reinhard, C. L. Remington, David C. Rentz, Chandler Robbins, Vincent D. Roth, E. P. Rouse, R. E. Ryckman, R. I. Sailer, Earl C. Schriver, G. G. E. Scudder, William M. Shields, David R. Smith, E. Graywood Smyth, Robert M. Stabler, Sarah H. Stabler, L. R. Steeves, George C. Steyskal, Frederic M. Stiner, Jr., Wallace A. Tarpley, Jack D. Tiner, Peter Tirrell, Charles A. Triplehorn, Neely Turner, William J. Turner, S. L. Tuxen, Nicolaas A. M. Verbeek, Kenneth A. Walker, Judith Stenger Weeden, R. L. Wenzel, A. Wetmore, Nixon Wilson, D. M. Wood, D. L. Wray, Russell E. Wright, Lawrence Zeleny, and F.

Introduction 9

Zumpt. Inasmuch as detailed records are not given except for type series and special cases, the individual contributions are ordinarily not elaborated. We humbly regret if any persons in the many years of this project have been overlooked.

History of the Classification of Bird Blow Flies

The genus Protocalliphora was proposed in North America by Hough (1899a) for Musca azurea Fallén (designated as type species) and M. chrysorrhoea Meigen. Both names were originally proposed for European species but then also used in North America. Hough unquestionably described a genus of bird blow flies, but his designation of azurea has led to arguments and problems. What is azurea? Does it belong to a different genus? Is it really Protophormia terraenovae (Robineau-Desvoidy)? Inasmuch as azurea is the type species of Protocalliphora, we must digress from the narrative to discuss its status even though we know now that the specific name does not apply to a Nearctic species. Relevant publications will be considered chronologically.

Fallén (1817): Musca azurea named and described, from "¢°," sent to him by Gyllenhal from Westergéthland (modern Vastergétland) Sweden. The number of specimens was not indicated.

Fallén (1821): M. azurea redescribed essentially as in 1817 but with a few additional details consistent with Protophormia terraenovae ("Squama nigricans," etc.).

Meigen (1826: 63): stated clearly that Fallén’s azurea included two species, and that only the male is azurea. The squamae (i.e., the calypteres) are described as white, and the description fits Protocalliphora as we know it. If he was dealing with original material, Meigen is the first reviser, in limiting azurea to males with white squamae. He visited Fallén at Lund and at least twice, in prefaces to his volumes, recorded his indebtedness to Fallén: "Die meisten von dem beriihmten Professor Fallen in Lund in Schweden 6ffentlich gemachten Arten, haben wir durch des Verfassers Freigebigkeit gleichfalls zur Ansicht und Vergleichung gehabt" (Meigen, 1818: xix); and Meigen, 1824: vi, in which he described his visit to Lund in 1823 in the company of the dipterist Wiedemann. They were received, Meigen stated, "with open arms" by Professors Fallén and Zetterstedt, who made their collections readily available for study by the visitors. Meigen undoubtedly saw original material, and his recognition of a mixed series is reinforced by Zetterstedt.

10‘ Protocalliphora

Zetterstedt (1838: 657) revised the Fallén material of azurea and recognized two species: azurea in the sense of a species of Protocalliphora and Musca groenlandica n.sp., now a synonym of Protophormia terraenovae. His description of azurea fits Protocalliphora for the most part, although he still says "squama nigricante." He specifically mentions Fallén’s material, "which that author had before his eyes when he was about to describe his species." Later he divided the collections and sent the Fallén Collection to Stockholm. Van Emden (1954) accepted Zetterstedt (1838) as first reviser.

Villeneuve (1918), under the combination Phormia azurea, noted that Zetterstedt’s description of Musca azurea was based on "le type méme de Fallén," and Stein (1924) referred to his study of "Die Type in Lund." Neither labeled any specimen of azurea as type or lectotype, nor gave any way of recognizing the specimen referred to.

Townsend (1931), who visited the collections at Stockholm and Lund, recorded the holotype in Stockholm, but no specimen there bears Townsend’s label. Curiously, he called the species Protocalliphora, even though the specimens in the Fallén Collection at Stockholm are Protophormia terraenovae (Ringdahl 1937, 1945; Hennig 1939; Sabrosky 1956).

Ringdahl (1937) stated that the two males and a female of Musca azurea in the Fallén Collection in Stockholm are identical with Phormia terraenovae Robineau-Desvoidy and groenlandic Zetterstedt.

Hennig (1939) borrowed a male of azurea from Stockholm, which he referred to as "der Typus." He dissected and figured the male genitalia, and recognized that the specimen belonged to the saprophagous genus Protophormia. He did not label the specimen as type or lectotype, perhaps assuming that the specimen sent him was the type. One of the two males in the Fallén Collection is obviously the specimen dissected by Hennig; it bears a printed label "hiervon micr. Prap./Kopulat.- Apparat," and a small pink label with a number that indicates it was loaned to Hennig.

Ringdahl (1945) also synonymized "the true Musca azurea" with Phormia groenlandica Zett., after "the type specimen in the Riksmuseum" in Stockholm. Again, he neither labeled nor identified a type or lectotype.

Sabrosky (1956) reviewed the problem of "The nomenclature of Protocalliphora," concluded that Meigen was really the first reviser, and stated that in his opinion “action of a first reviser takes precedence over later actions, even if one of the latter

History of the Classification of Bird Blow Flies 11

involves selection of a lectotype or neotype." He then designated a lectotype consistent with Meigen’s revision, choosing an old unlabeled male in the collection at Lund.

With the advantage of hindsight and the subsequently developed International Code of Zoological Nomenclature (1961, 1985), we now conclude as follows:

1. Lectotype designation has been given precedence over restriction by revisers (ICZN, Art. 74a.ii, 3rd edition; cf. also Rec. 74A). Sabrosky’s 1956 point of view in favor of reviser action did not prevail. Unfortunately, as a result, a heedless or inadvertent designation of lectotype can upset usage long established by revisers.

2. The material sorted by Zetterstedt contained two species, one a Protocalliphora, the other a Protophormia. From the labels, most of the former now in the collection at Lund were collected after Musca azurea was published and therefore cannot possibly be the syntypes, or part of the original series. One specimen, unlabeled as very old specimens often are, might have been an original specimen and at least cannot be demonstrated not to have been. This was the specimen selected as lectotype by Sabrosky (1956), who then believed that the first reviser (Meigen 1826) limited the species to a specimen of Protocalliphora. The other specimens that also appear to be original material, two males and one female, are in Stockholm; these are Protophormia terraenovae.

3. Once the first reviser approach is rejected, then the question of lectotype must be reexamined critically. Did any of those who referred to "the type" or its equivalent in any language actually designate a lectotype? None of those so labeled a specimen, but labeling is not specifically required by the Code. If not labeled the specimen must surely be described in some way so that it can be recognized as the type; otherwise the so-called designation is ambiguous and imprecise and cannot be regarded as valid. Hennig, who dissected the male genitalia, appears to qualify because the specimen he studied is labeled as dissected, and it also bears a numbered pink slip of paper that indicates it was loaned to Hennig. True, Hennig did not see all of the original material and in a sense did not really revise, but under Article 74b of the Code his inference that the specimen he studied was "the type" is deemed to have been lectotype designation should other syntypes be discovered. This assumes, of course, that the lectotype was one of the original syntypes, and it appears to have been, or at least cannot be demonstrated not to have been.

12 — Protocalliphora

4. We are thus led to the unfortunate conclusion that Hennig’s (1939) figured specimen is the lectotype, and that accordingly Musca azurea in the strict sense is a Protophormia, with priority over the relatively common and widespread species long known as Protophormia terraenovae (Robineau-Desvoidy).

5. All authors are agreed on preserving the generic name Protocalliphora on the basis of misidentified type species (ICZN, Article 70), because Hough’s description clearly applies to the bird blow flies and the name has always been used in that way. Peus (1960) agreed that the type species was misidentified, but considered that Hough based his genus on American species then identified incorrectly under the European names. There is nothing in Hough’s paper, however, to suggest that he saw only American examples. Furthermore, there is evidence in a paper he published the same year (Hough 1899b) that European material was available to him ("I have compared my American with European specimens from Prof. G. Strobl and Dr. O. Schmiedeknecht"). We can therefore accept his designation of azurea Fallén as referring to a European species.

6. The question of the correct specific name for azurea sensu Hough and European authors is not so easily solved. The next oldest name that has been associated with the bird blow flies is Phormia caerulea Robineau-Desvoidy, 1830, but no type material is known to exist and the description is not identifiable. In view of the extensive usage of the name azurea Fallén for the bird blow flies, Protocalliphora, we believe the best course would be to suspend the rules and fix the name azurea for a species of those flies, thus also leaving the widely used name Protophormia terraenovae undisturbed.

Sabrosky (1984) applied to the International Commission on Zoological Nomenclature for an amendment to the International Code that would relax the present requirement that cases of misidentified type species must be referred to the Commission. In actual practice many taxonomists have proceeded unilaterally to recognize the species actually before the original author, when the circumstances are considered clearcut that it was misidentified. Although no official decision has yet been published, we understand that reaction was favorable. We therefore continue to use the name Protocalliphora for the bird blow flies in the belief that ultimately this name will be approved, whatever the mode of achieving it. Meanwhile a formal application to the Commission has been prepared (Sabrosky 1988), because it is still worthwhile to fix the use of azurea for a bird blow fly and thus to avoid the confusing

History of the Classification of Bird Blow Flies 13

transfer that would substitute azurea for the widely used name terraenovae in Protophormia.

Hendel (1901) erected a new genus, Avihospita, for azurea and relatives, but Aldrich (1901) promptly pointed out that it was a synonym of Protocalliphora Hough, with the same type species. The genus Protocalliphora was recognized as distinct in both the Nearctic (Aldrich 1905) and Palearctic catalogues (Bezzi and Stein 1907). On the other hand, one of the most respected of European specialists on muscoid Diptera, Villeneuve (1911), rejected the genus as useless and pronounced it a synonym of Phormia. In later publications (1928, 1931) he referred to it as at best no more than a subgenus. The genus has usually been maintained as distinct by American dipterists, however, as in the well-known manuals by Williston, Curran, and Townsend, the revisions of Nearctic Calliphoridae by Shannon (1923, 1926) and Hall (1948), and in the new Manual of Nearctic Diptera (Shewell 1987), and it is recognized as distinct by modern European dipterists. The distinctive biology and larval morphology are important factors in its recognition.

Aside from occasional records, there was little attention to the genus in North America in the first two decades after its proposal, and it was regarded as rather uncommon, probably because specimens are seldom taken by ordinary net collecting. Harbeck (1907), probably based on an identification by Coquillett, apparently first referred to Protocalliphora a described Nearctic species, Calliphora splendida Macquart. Later, Townsend (1919) described Phormia metallica in this group, using Phormia instead of Protocalliphora because of a nomenclatural confusion.

The first revision of the North American species was by Shannon and Dobroscky (1924). The Palearctic names azurea and chrysorrhoea were dropped and three Nearctic species were recognized, avium and hirudo (now braueri) as new species, and splendida (Macquart) (syn., metallica Townsend), together with one variety of avium, three varieties and a subspecies of splendida, and one variety and one subspecies of hirudo. The male terminalia were utilized to some extent, but except in avium with its unusually broad surstyli (outer forceps) the differences were not fully appreciated and were dismissed as being too intangible for the differentiation of species.

The second revision of the Nearctic species was that of Hall (1948) in his book on The Blowflies of North America. Hall recognized ten species, raising four of Shannon and Dobroscky’s "varieties" to specific rank and describing three new species. He

14 ~~‘ Protocalliphora

also proposed a new genus, Apaulina, for the Nearctic species, based in part on certain differences from the Palearctic species pointed out by Shannon (1923) and by Shannon and Dobroscky (1924). Apaulina is considered by us to be a synonym of Protocalliphora (cf. later discussion), and the synonymy has already been published by several authors on the authority of Sabrosky in litt.

The present revision, the third for North America in 60 years, includes a total of 26 species, including 15 described as new and two (chrysorrhoea and braueri) recognized as Holarctic. True chrysorrhoea is here recorded in North America for the first time, but braueri was previously known as a Nearctic species, hirudo. Some additional species appear to be new, but these are left undescribed because of inadequate material. A general discussion of this taxonomic revision and of the field and laboratory work was presented both as an invitational paper and a demonstration at the Tenth International Congress of Entomology at Montreal in 1956 (abstract, Sabrosky and Bennett 1958), and as a contributed paper at the Thirteenth Congress at Moscow in 1968 (short abstract, Sabrosky and Bennett 1971).

Evidence from field and laboratory studies of both adults and immature stages supports the view that the genus is composed of a number of distinct species albeit closely related and confusingly similar, and not of a few variable species with color forms or ecological races. Additional evidence has been furnished by a study carried out by Prof. J. W. Boyes of McGill University, who found good specific differences in the somatic chromosomes of six eastern species, aenea, avium, hirundo, metallica, shannoni (as n. sp. near sialia), and sialia, made available to him at Algonquin Park, Ontario (Boyes 1961, Boyes and Brink 1965, Boyes and Shewell 1975).

Geographic Distribution

The genus Protocalliphora is Holarctic and predominantly northern (Maps 1, 2). In North America it occurs in Alaska, throughout Canada, Greenland, in most of the contiguous states of the United States, and in the Nearctic part of Mexico. The flies seem to occur chiefly at higher altitudes in the southern extensions of the range. There are only a few scattered records from the southeastern states, the southern Mississippi Valley, and throughout the Great Plains states and provinces. In the East, specimens have been collected in the Appalachian Mts. south to northern Georgia. The southernmost eastern record, from south central Georgia (Norris 1958), is based on a single puparium

Geographic Distribution 15

found in the nest of a brown-headed nuthatch. In the West, specimens are known from California as far south as San Diego and from Baja California Norte, from various localities in Arizona and New Mexico, especially in mountain areas, and from six localities on the Mexican Plateau (the southernmost: in Morelos, Puebla, and Tlaxcala states south and southeast of Mexico City). The few Mexican localities, with a total of 11 net-collected specimens, highlight the extensive Nearctic territory in Mexico that is terra incognita for Protocalliphora.

It remains to be seen whether the absence of records in some states reflects merely lack of collecting. We have made numerous efforts to find material from some of those states, and to encourage collecting of birds’ nests in a _ search for Protocalliphora, but thus far with little or no success. Lack of collecting would be especially noticeable in a group like Protocalliphora, which requires rearing of immature stages rather than ordinary net collecting of adult flies.

The map (Map 1) of course does not necessarily show the abundance of Protocalliphora. A dot on the map may represent only a single net-collected individual, or only one species, or it may represent hundreds of specimens of several species, or repeated recoveries over a period of years. The major areas of collecting (marked with large dots), in terms both of number of specimens and of species, are Kamloops, B.C. (G. J. Spencer and others), Ravalli County, Mont. (W. L. Jellison and C. B. Philip), northern Utah (T. L. Whitworth), Algonquin Park, Ont. (G. F. Bennett), Ithaca, N.Y. (R. C. Shannon, I. D. Dobroscky and others), eastern Massachusetts (many localities close together, the records published by C. W. Johnson 1925-32), and California localities (university staffs and students).

In the Palearctic Region (Map 2) the genus also appears widespread, with numerous records from Europe (not individually spotted for present purposes), but only widely scattered records from temperate Asia. North Africa and vast stretches of Palearctic Asia are virtually bare of records. There are published records from Algeria, from the Altai Mountains in central Asia, from the Ussuri Region near eastern Manchuria, and from Japan. There are hitherto unpublished records in the collection of the National Museum of Natural History from Korea, China (Shanghai, and Harbin in Manchuria), and from the Wa-Hu Pass, 16,400 ft., on the border of Sichuan (Szechuan) and Tibet. Protocalliphora azurea was recorded from Iceland in 1889 and 1890, but the records are now doubted and the species has not since been recovered (Nielsen et al. 1954). This may reflect

16 ~—-Protocalliphora

lack of collecting from birds’ nests. For a half century Protocalliphora was known from Greenland from a single specimen collected in 1892, and the next specimens (reared) did not turn up until 1952 (cf. P. tundrae).

Any general statements must of course be qualified by "as far as known." The conspicuous lack of records from some areas and regions (Map 1) often reflects lack of collecting, or lack of rearing from birds’ nests. Net collections yield very few specimens, and the full extent of a fauna can only be determined by the time-consuming method of locating birds’ nests and rearing any mature larvae or puparia found after the young have left the nest. Unverified published records of species cannot usually be relied upon in view of the complexity now known to exist in the genus, although identifications by competent dipterists can be accepted as showing the presence of the genus Protocalliphora.

One odd record should be noted, only to dispose of it: Protocalliphora azurea (Fallén) from Kona, Hawaii (Grimshaw 1901: 27, as Calliphora azurea ). Both Adrian C. Pont of the British Museum (Nat. Hist.) and Sabrosky have examined the specimen and find that it is actually the black blow fly, Phormia regina (Meigen). The genus Protocalliphora has never been found in the Hawaiian Islands. Hardy (1981) was in error in assigning this specimen to Chrysomya megacephala (Fabricius).

The Palearctic Fauna

Before dealing with the taxonomy of the Nearctic species, a brief review of the Palearctic fauna will be useful for comparison. Although the generic distinctness of the bird blow flies was first recognized by an American, some knowledge of the Palearctic species long antedated that of the Nearctic fauna. In the last few decades, knowledge of Protocalliphora in the Palearctic Region has expanded much like that in the Nearctic Region. For many years, European dipterists recognized only one or two species, azurea Fallén (sordida Zetterstedt) and chrysorrhoea Meigen, the latter often called a variety or subspecies of azurea. In the older literature, usage of azurea has been so varied, and so many species have been described since azurea was used in a broad sense, that it would seem hopeless to straighten out the published literature without a thoroughgoing revision based on good reared series of both sexes supported by larvae and/or puparia, followed by reexamination of all existing types and of voucher specimens for published records.

17

The Palearctic Fauna

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18 Protocalliphora

Table II, end Specific names of uncertain or disputed identity

** 1780. reviso Harris (Musca). Questioned as synonym of Protocalliphora azurea by Pont 1976.

** 1829. carnarida Stephens (Musca). Nomen nudum, synonym of P. sordida by Pont 1976.

1830. caerulea Robineau-Desvoidy (Phormia). Senior synonym of Protocalliphora sordida by Séguy 1929, but synonym of P. azurea by Zumpt 1956, 1965, and Schumann 1986.

* 1838. violacea Meigen (Musca). Listed by Rodhain & Bequaert 1916, from Meigen’s type, as male of Protocalliphora sordida Zett., and by Schumann 1986 as a nomen dubium in Calliphorinae. Villeneuve wrote to Aldrich on Feb. 2, 1923 (card in U.S. National Museum of Natural History, Washington, D.C.) that the male type is undoubtedly Phormia (Protocalliphora) azurea(Fallén), confirmed by Knut Rognes, n. syn. in litt., but without dissection of the type. Whatever its true identity, the name violacea cannot be used in any case because of primary homonymy; it is preoccupied several times over in Musca, by Scopoli 1763, Panzer 1804, and Fabricius 1805.

1863. nigripalpis Robineau-Desvoidy (Phormia). Synonym of Phormia caerulea by Bezzi & Stein 1907, but synonym of Protophormia terraenovae (Robineau-Desvoidy) by Hall 1948, Zumpt 1956, 1965 (both as nigripalpus), and Schumann 1986.

* 1863. corusca Robineau-Desvoidy (Phormia). Synonym of Phormia caerulea by Bezzi & Stein 1907, but of Protophormia terraenovae by Schumann 1986.

* 1867. nidicola van Heyden in Nowicki (Calliphora). Nomen nudum, synonym of Protocalliphora azurea in Bezzi & Stein 1907, but of Trypocalliphora braueri in Schumann 1986.

* = Not mentioned by Zumpt 1956. ** — Not mentioned by Schumann 1986, nor Zumpt 1956.

The Palearctic Fauna 19

Although a few of the recent species have been based on series of specimens, in general most species were described and named on the basis of very inadequate material (cf. Table II). Of the 22 available names (15 presently recognized species and seven synonyms; Schumann 1986), the type material of only seven was reared, including one in part. For the older species, the number of specimens was not specified, but in most cases was probably only one or two, judging from the surviving material in the well-preserved Fallén, Meigen, and Zetterstedt collections. The exception is Lucilia dispar Dufour, of which at least 80 were reared. Most disturbing, eight of the 14 Protocalliphora described in and since 1956 were based on a total of 14 specimens (11 males, 3 females, with only four of the 14 reared), and seven were described from one sex only.

Three names were first proposed as subspecies or varieties, in addition to the frequent treatment of chrysorrhoea as a subspecies of azurea. In three of the four cases, our experience in the Nearctic fauna convinces us that the characters given mark distinct species. Recognizing these as good species, and accepting some published synonymy--although with realization that a broadly based revision is needed, the presently known Palearctic species of Protocalliphora (including Trypocalliphora, see later discussion) can be listed as in Table II (species listed chronologically to show the historical sequence of the development of knowledge). :

The revisions by Gregor and Povolny (1959) and by Peus (1960) contain keys to the adults of the Palearctic species and figures of male and female terminalia. Other figures of male terminalia were published by Séguy (1929), Hennig (1939), Zumpt (1956), Kano and Shinonaga (1966), Grunin (1966), and Rognes (1985). Figures of the immature stages were published by Rohdendorf (1957) and Ishijima (1967), although the immature stages are neither described nor figured in the detail of those in the present paper. The cited publications together with available Palearctic material of adults of nine species and puparia of five have made possible comparisons of the Palearctic and Nearctic species. Two species are recognized here as Holarctic, one, chrysorrhoea, virtually restricted to nests of a Holarctic bird, the sand martin (of European literature) or bank swallow, Riparia riparia (L.), in a distinctive ecological niche. Another species, braueri (Syn., hirudo), is now recognized as Holarctic (Rognes 1985), although Peus (1960) had a different opinion and described the European form as a distinct species, Trypocalliphora lindneri (see discussion under braueri).

20 = Protocalliphora

The literature on Palearctic species is usually not relevant to the present study and it has been referred to sparingly. It has been summarized by various authors, including the revisions cited in the preceding paragraph and the authors cited in the second paragraph of the introduction. There are numerous host records, but the Palearctic species of Protocalliphora have not been studied with the detailed ecological approach reported in the present study. It seems to us quite probable that more intensive biological and ecological studies, and attention to the characters of the immature stages, at least of puparia or 3rd-instar larvae, might reveal additional species in the extended Palearctic Region. Noteworthy also is the dearth of information about some of the species already named but known from single or few net-caught specimens, and association of sexes might even be questioned in some cases. The immature stages are unknown or undescribed for most of the species. These are difficulties that we are in no position to solve, but as far as possible we have compared the Nearctic and Palearctic species before publishing our new species.

Life History

The entire life history has not been followed for a single species of Protocalliphora in North America, and perhaps not anywhere in the world, but enough is known of parts of the cycle and the factors affecting it that inferences can be drawn and reasonable statements made. Observations and experiments were recorded by Bennett (1957), Whitworth (1976), and Gold and Dahlsten (1984).

Species of Protocalliphora apparently overwinter as adult flies. Overwintering in the egg or larval stages would be impracticable in most cases, even if they were cold tolerant, inasmuch as most birds do not utilize old nests, and such nests exposed to the weather are usually destroyed by the following season. Overwintering in the pupal stage would theoretically be possible, but evidence from Bennett’s studies suggests that the pupae of these flies are also not cold hardy. Other evidence also points to the adult stage: (1) Adult flies emerge in mid or late summer from puparia in the nests of the last brood of birds although no more hosts will be available that season; (2) the length of life of adults can be long; (3) only empty, dead, or parasitized puparia are found in old nests taken in fall or winter (e.g., Dobroscky 1925); and (4) there are numerous late and early season records of adult flies, and even a few winter records.

Life History 21

Johnson (1930) listed a number of fall, winter, and spring captures of live adult Protocalliphora as indicating that "there is no doubt that Protocalliphora hibernates as an adult and awaits the arrival of the birds in the spring to oviposit in their nests." The winter dates of live specimens are especially significant. One specimen, a male P. bicolor, was collected at Rumney, N.H., on December 22 by P. J. Darlington, Jr., in chopping up a dead pine. Another specimen, a female P. shannoni, was collected at Concord, Mass., Jan. 10, 1925. Stiner (1969) recorded a frayed-winged male P. metallica found under bark on April 22, 1968 at Newark, Del., and from abundant evidence on nesting songbirds in the area he concluded that it could not have developed in a 1968 nest. From the timing plus its "old" appearance, the individual appeared to have overwintered. In Norway, Somme (1961) recorded finding live P. azurea in crevices in mountain huts at 900 and 950 meters in March and early April. Adults of Protocalliphora can be long lived, as shown by Bennett’s studies on their longevity, in which four species averaged 75-83 days, with maxima of 170-281 days. Whitworth (1976) found some adults still alive after eight months, in a cage exposed to winter weather but with sawdust offering available protection for the flies.

The available records indicate overwintering by both sexes, rather than by fertilized females alone as in other Calliphoridae known to overwinter as adults--e.g., the related Phormia regina (Meigen) and Protophormia terraenovae (Robineau-Desvoidy). Sperm apparently remain viable for long periods. Bennett found that sperm removed from the seminal receptacles of Protocalliphora avium 119 days after the last insemination appeared to be as motile as those removed only 10 days after insemination. Overwintering females might still be fertilized, however, and overwintering males only occasional survivors or infertile or ineffective. It would seem a more efficient adaptation in the life history of Protocalliphora for overwintering females to be fertile and ready to oviposit at once in the nests of the first broods of the early-nesting birds. An especially relevant bit of evidence here was contributed by the late G. J. Spencer (personal communication), who informed us that live puparia of Protocalliphora were found in magpie nests in British Columbia in February, the magpies nesting when snow was still on the ground. Obviously, gravid adult females were present in the winter and became active enough on warm days to deposit eggs in the new nests. It is of course still possible, with both sexes

22 ~~ Protocalliphora

overwintering, that mating could occur when adults become active on warm days.

There is some evidence that males of Protocalliphora may gather at high, bare aggregation sites as do some other muscoid Diptera. Dodge and Seago (1954) recorded two "spp.", now known to be sialia and braueri (hirudo), on mountaintops in northern Georgia. For P. spatulata, records from Montana, Wyoming, and New Mexico specify the top of a peak or of a range. A number of other specimens are labeled from specific mountain peaks, and some of these may also have meant the mountaintop.

Female flies apparently oviposit in or on nests containing very young nestling birds. Although Zumpt (1965) stated that it is not known whether Protocalliphora is oviparous or larviparous, numerous observations now show that these flies are oviparous. In those studies, of course, the nesting birds were usually not disturbed, and in order to find mature larvae and puparia of Protocalliphora the nests were not collected until after the young had fledged. Bennett (1957) and Whitworth (1976) had numerous records and experiments with eggs deposited by females caged in the laboratory. In Europe there have also been studies on the Oviposition of P. azurea on the great tit, Parus major L. (Eshuis-van der Voet 1972, 1975, and with others, especially Eshuis-van der Voet and Kluyver 1971, who established that the eggs were laid "in the nest-material, close to the nestlings.").

A unique observation is that of Meng (1954), recorded in an obscure and hard-to-find paper that summarized observations on P. avium during an intensive 4-year study of the Cooper’s hawk in New York State, that "as soon as the hawk eggs start hatching, the adult flies, which look very much like bluebottle flies, deposit their eggs along the edges of the nests. Soon after being laid the fly eggs hatch, and the larvae find their way into the ear openings of the young hawks." Professor Meng (personal communication) has assured us that on several occasions he saw the flies laying eggs. A suggestive supporting observation is that of Law (1929), who observed a large fly looking like Protocalliphora, which, after some circling, finally entered a nest of the pygmy nuthatch, Sitta pygmaea (San Bernardino Mts., Calif.). Law further stated that "On many occasions I have seen these blue flies circling the entrance to nests containing the young of nuthatches, chickadees, and bluebirds."

Rausch (1972), in discussing cutaneous myiasis by P. hirudo (= braueri) in the head of a fledgling Wilson’s warbler, speculated that the aggregation of larvae in a single locus in the head

Life History 23

indicated the likelihood that "the eggs are deposited directly upon the head." Rognes (1985), in reviewing the literature and records on Trypocalliphora braueri, chiefly in Europe, concluded that the eggs were "probably laid in clusters directly upon nestlings of various ages, apparently also on newly hatched ones." Whitworth (1976), who often observed adults of chrysorrhoea about bank swallow burrows, recorded that "on one occasion, an adult was observed in a nest, walking from one nestling to another while dipping its abdomen. Eggs were laid on several nestlings in small clumps and were attached to the feathers. No eggs were found deposited in the nest material." On the other hand, the observation of Meng (1954) suggests that even in the case of burrowing larvae, eggs might be laid on the nest and the young larvae could find their own way to an appropriate feeding site.

Tirrell (1978) reported that females of P. avium "apparently laid eggs directly" on nestling red-tailed hawks. He observed flies "entering the ears of two nestlings from which larvae had been removed. Subsequently, each ear cavity was infested within one day by approximately 40 uniformly sized larvae, each 2.5 mm in length." In earlier correspondence (Feb. 17, 1976), Tirrell had elaborated on his field notes during hours of observations: "Typically, there were 5-20 adult flies around the nest. One or two flies continuously attempted to settle on a nestling’s head and to walk to an aural cavity. Their success per attempt was very low, but at least several times an hour, a fly attained the edge of an aural cavity. These flies usually moved their heads in and out of the ear opening, then turned and held the posterior near the edge of the cavity, or walked around the edge, stopping frequently, and touched the end of the abdomen on the edge." Although eggs were not then observed, he "interpreted this as egg-laying behavior," especially when re-infestation occurred so quickly in cases where he removed all the larvae and reexamined within a day. Also, on several occasions he found scattered clumps of eggs near the axillary areas of the nestlings, showing that indeed eggs had been laid on the body.

The egg stage is short, probably 24-48 hours, although this is an inference because of uncertainty about the exact time of Oviposition. In the laboratory, Bennett found that eggs laid by P. sialia hatched within 24 hours, but Whitworth (1976) recorded 38-43 hours for P. chrysorrhoea and 72 hours for P. asiovora. In nature, eggs laid on the nests of smaller birds with short nestling period would necessarily have had to hatch quickly for larval development to be completed before the young birds fledged and left the nest.

24 Protocalliphora

Details of the life history and biology will be given in a later publication. There are some differences in the data from the respective studies, probably affected by differences in temperature, the species of hosts, and the species of flies, and also affected by uncertainty of the time of oviposition and the duration of the egg stage and the larval instars, with some inferences necessary. The time in the Ist and 2nd larval instars is relatively short, with a longer period for the 3rd instar, the three totaling 7-15 days. The prepupal period is short, varying from 1-4 days, and the puparial period has a wide range, 9-36 days or perhaps even more.

There may be very few larvae in a nest, especially on small birds, or there may be surprisingly large numbers. Séguy (1955), in a general review of Protocalliphora, no doubt chiefly from the European literature, stated that "one finds from one to thirty larvae per nest," but that is an understatement in our experience. In contrast, Bennett found 970 in one nest (Sabrosky and Bennett 1958), with other records of 400-800 per nest, and Whitworth recorded larval populations of 300-1200 per nest. In general, the numbers of larvae and/or puparia reported in the literature are much lower, although occasionally in the hundreds (e.g., Dobroscky 1925, 343 ina nest of an American crow; Mason 1944, 442 in a nest of an American tree swallow; Kenaga 1961, 206 in a nest of an eastern bluebird; and Gold and Dahlsten 1984, 273 in a nest of a chestnut-backed chickadee). Whitworth (1976) has given a detailed discussion of a number of factors that affect larval populations in nests. Large numbers of larvae may result primarily from large nests with suitable hiding places rather than from large birds. Whatever the bird size, small or fragile nests are inadequate shelter for very many larvae.

Various authors have observed that the larvae are "tough". One dramatic bit of evidence was provided by Plath (1919a, b). He had placed some larvae in a fixing fluid for six hours, washed them in 50% alcohol, and placed them in 90% alcohol, but two days later they were still alive! Moreover some remained alive for two or three days in "very strong insect powder," which makes one wonder at some of the control recommendations that have been made.

The larvae of Protocalliphora are obligatory blood-sucking parasites. Bennett closely observed feeding by the larvae, and observations were also made by Whitworth (1976). Bennett further details the feeding as follows: The anterior part of the body, with mouth hooks thrust out, turns like a person turning a paring knife, with a cutting motion. It penetrates the skin a

Life History 25

little way, then thrusts in the prothoracic fringe, which points backward and apparently holds the maggot in the wound. Then the maggot cuts farther in with the same cutting motion, followed by another grip with the fringe. At first the maggot braces itself against the nest wall, sticks, or other material. When it is well into the skin it relaxes, and starts to suck blood. It takes a long time for this, about 45 minutes to get the crop one-third full. The amount of blood can be seen through the body wall. Fully-fed larvae are distended with bright red blood, which gradually darkens and decreases in size as digestion and excretion proceed. Some larvae burrow into the skin and may embed themselves (P. braueri, apparently typically), but even these larvae are basically blood-sucking. Rognes (1985), discussing braueri in Europe, stated that "the digestive tract of the larvae mostly contains blood," even though the larvae "make permanent subcutaneous burrows" and "extensive tissue destruction may occur." Arnold (1919) had recognized this long ago, pointing out that "the maggot eats a burrow or chimney into the flesh of its victim, remaining stationary and feeding upon the fluids." Although larvae of braueri are often found attached to their host, some larvae containing a blood meal have also been found by Bennett in nest material.

Third-instar larvae take two or three blood meals for maturity to normal-sized individuals. Apparently they can complete development on only one blood meal, but the resulting adults are undersized, or "runts."

Raptorial birds in particular seem characteristically to have Protocalliphora larvae in the ear cavities, and numerous authors have recorded this (e.g., Burtch 1920, Sargent 1938, Hill and Work 1947, Hamerstrom and Hamerstrom 1954, Meng 1954, Hickey 1969, Tirrell 1978). The Hamerstroms and Meng, who have studied myiasis in hawks in considerable detail, differ on whether or not the larvae remain attached until mature. The Hamerstroms believed that the larvae "spend the entire time between hatching from the egg until pupation within the hawk’s ear," whereas Meng stated flatly that "the maggots are intermittent feeders and do not remain in the ears continuously." Certainly the ears are not large enough to contain fully fed larvae in the numbers that have been reported. The Hamerstroms have graphically described what can happen: "We doubt whether it makes much difference how many maggots there are in the ear. We have never seen just a few; there have always been either a goodly number or none at all. As the young maggots grow larger, they become crowded in the limited space of the ear cavity.

26 ~~ Protocalliphora

Breathing space is probably even more at a premium than feeding space. One by one the surplus maggots dropped out of the ears of our Cooper’s Hawk’s and fell to the floor of the cage where they died, until at last each ear contained what looked like the maximum number of maggots that could fit into the distended opening. These did not leave the ear prematurely and pupated successfully." In nature, surplus maggots would probably have dropped into the nest material and survived to feed elsewhere on the body.

A few of the other published records are detailed enough to bear repeating. Meng (1954) wrote that "When the [Cooper’s] hawks are very small the only indications one has that they are infected are the droplets of dried blood around the ear openings. The larvae grow rapidly on their diet of blood, and by the time the eyases are three weeks old their ear opening may be completely plugged by the protruding ends of the screw-worm fly larvae" [meaning in this case the bird nest screw-worm, P. avium]. Hill and Work (1947) found two young sparrow hawks [i.e., American kestrels] with a black crust around the openings of the ears: "When this had been removed, it became apparent that the ear canal was markedly swollen and filled with fat, squirming, grayish larvae. The right nostril and both ears of each bird contained larvae; none was found elsewhere. The nostrils were deformed, as the diameter of the larvae found there far exceeded the width of the normal nasal passages."

Do Protocalliphora Larvae Kill Young Birds?

This is a challenging question, often asked, and of special interest to the many people interested in birds and their welfare. A flat yes or no answer is impossible. Opinion is divided, probably depending on the particular circumstances or the experiences of each author. In certain cases, death of the nestlings can clearly be ascribed to the maggots of Protocalliphora, as when larvae in the head penetrate to the brain. In other cases, however, nestlings seemed to be relatively unaffected in any way noticed by the observers. There is always the possibility that the larvae contributed to stress or weakness that resulted in death later from other causes, such as unfavorable weather, malnutrition in periods of food shortage, and other parasites or diseases such as arborviruses, Plasmodium, Leucocytozoon, or Haemoproteus. For example, Wing (1956:405) in his book on the Natural History of Birds concluded that "In

Protocalliphora Larvae = 27

inclement weather when little food may be brought to the young, the attacks of the parasites upon weakened nestlings may cause death." One can understand that this could happen especially in the case of heavy infestations, hundreds of larvae, on small birds. Some authors have also recorded nestlings smaller or weaker than normal, and a longer than normal nestling period, in the presence of large numbers of maggots. Whitworth (1976) has pointed out the likelihood that birds losing blood daily to Protocalliphora "would be somewhat anemic and perhaps less able to withstand additional stresses."

It is our impression that the literature contains much anecdotal material on which assumptions have been based, rather than factual evidence. Statements seem to reflect a belief that because the larvae are hematophagous, they must ipso facto be considered lethal or at least potentially life threatening. In many cases, the feeding of Protocalliphora larvae may be no more serious to the host than the feeding and irritation of fleas on a dog. On the other hand, comments in the literature as to little or no evident injury might have reflected slight or relatively low numbers of maggots, or large numbers of maggots but in times of favorable weather and abundant food when the nestlings were vigorous.

Protocalliphora may also be, on occasion, unjustly accused of causing the death of nestling birds. In at least one case that we have been able to investigate (Munro 1949), the presumed Protocalliphora larvae taken from dead fledglings proved, from reared adults, to have been the scavenger calliphorids Phaenicia sericata (Meigen) and Phormia regina (Meigen), which had invaded the carcasses subsequent to their death from other causes (or regina might have been directly involved in myiasis). This is further borne out by the observation of Coutant (1915), who investigated the death of a nestling American crow infested by Protocalliphora larvae and found that the crow actually died of a "malarial fever" and that the erythrocytes contained "enormous quantities of Halteridium" [=Haemoproteus, protozoan blood parasites]. In another case, "Protocalliphora" larvae submitted for identification as having killed barn swallows, undermining the back skin and "clear down next to the ribs in a bloody necrotic mess in one bird," proved to be larvae of Phormia regina.

Some authors have recorded dead birds along with infestation by Protocalliphora, although not assigning definite responsibility for the mortality, appearing to leave guilt by association. For example, Shannon and Dobroscky (1924) listed the type series of hirundo as 64 specimens from cliff swallow nests, adding "Many

28 — Protocalliphora

nests examined had dead remains of young." Of material received for identification, two larvae and 12 puparia were found at Auburn, Ala., in the nest of a wren "where young birds failed to mature."

Eliminating pure speculation and mere repetition of the views of others, we will summarize the expressed opinions and evidence, with appropriate quotations.

Little or no injury to the nestlings was reported by a number of authors, with various comments, e.g., Jellison and Philip (1933) and Jellison (1949), no fledgling crows or magpies ever found dead in the nest; Stoner (1936), bank swallows "suffering no serious handicap"; Krug (1941), in bluebirds and tree swallows "the mortality is almost negligible"; Zeleny (1970), bluebirds are "usually able to survive," and blow fly infestation, unless heavy, "does no noticeable harm to the birds"; and Seidensticker and Reynolds (1971), "we would attribute no mortality [of hawks] to these larvae." Many authors expressed the caveat, however, that even though they observed no injury or mortality, heavy infestations of larvae might have those effects, or (Gold and Dahlsten 1984) might so affect the young that they fledged "in a weakened condition." McAtee (1929) stated unequivocally, however, that "heavy infestations [of Protocalliphora in bird houses] are not especially destructive to nestling birds." Whitworth (1976), while acknowledging that high numbers of Protocalliphora \arvae might be lethal, concluded from his studies of larval populations that "near-lethal numbers are uncommon under natural conditions." Bennett (1957) concluded that "in general it appeared that the Protocalliphora did not seriously harm their avian hosts," and Gold and Dahlsten (1984) found nestling mortality of chickadees "negligible," although they hypothesized that heavy infestations would result in weakened fledglings.

Except for references to braueri, discussed later, many European authors have not regarded Protocalliphora as unusually serious. Owen (1954) in a review in British Birds concluded that "it would appear that ... mortality of the host is exceptional and possibly always linked with other factors." A year later, Owen and Ash (1955) published additional British records of Protocalliphora infestation and again suggested "that any resultant mortality among nestlings is most exceptional." On the other hand, Boyd (1935, 1936) concluded that prevalence of Protocalliphora larvae "may materially affect the size of the broods and account for the broods of one or two young that are

Protocalliphora Larvee 29

often reported" and further that the larvae "probably cause many deaths among the young [barn] swallows."

Death or serious injury has been reported by a number of authors, although it is not always clear what is observation and what is presumption based on what others have written or on the large numbers of larvae found in nests. Sometimes the presence of larvae is linked with other factors as contributory causes of death. For example, Allen and Nice (1952) on purple martins reported heavy infestations of Protocalliphora larvae coincident with cold wet weather and concomitant shortage of food, resulting in the death of many nestlings. Zeleny (1970) found that while in general Protocalliphora infestation does no noticeable harm to young bluebirds, the blood loss from heavy infestations "seriously weakened" young birds and "may be a contributory cause to the death" for both nestlings and fledglings, and "Very heavy infestations ... may kill the young birds outright."

Infestation of the ears of birds by Protocalliphora larvae is more obvious than the somewhat hidden feeding on the under side of the body, and when severe it attracts special attention. Sargent (1938) reported so many maggots in the ears of a red-tailed hawk that the ear openings were stretched to twice normal size and "completely plugged by the caudal ends of the maggots." Tirrell (1978) and the Hamerstroms (1954) both noted that infested hawks shook their heads intermittently or scratched them, apparently irritated by the larvae, although apparently there was no permanent damage. Hagar (1969, on peregrine falcons), Crocoll and Parker (1981, on broad-winged hawks), and Bortolotti (1985, on bald eagles) found, respectively, "no apparent ill effect," "no major deleterious effects," and "no indication of any ill effects" from Protocalliphora infestations. Meng (1954) stated that none of the observed Cooper’s hawk eyases "seemed to be weakened appreciably." On the other hand, Bent (1937) recorded the results by a correspondent of skinning and dissection of two young red-shouldered hawks, in which the maggots had not only disfigured the outer ear cavities but had destroyed the ear drums. We conclude that, except for the occasional instance when such serious damage has been done, in general the large young of raptors can support sizeable numbers of maggots, the young birds have long nestling periods, the maggots mature and leave the birds, and the birds recover from any ill effects and fledge successfully. As the Hamerstroms remarked, when larvae left the ear, "By then our hawks were as large as branchers and made a good recovery."

30 ~—- Protocalliphora

Whitworth observed larvae commonly in the noses, ears, and inside feather sheaths of nestlings of its common hosts, especially raptors and corvids (crows, ravens, magpies). Whitworth (1976) noted that "the presence of these larvae is often indicated only by the scablike accumulations of larval excreta around cavity openings."

Subcutaneous infestation by Protocalliphora braueri(P. hirudo of Nearctic literature) is another feature of Protocalliphora that deserves special mention. A few authors have recorded serious damage by larvae of this species. R. C. Miller (1929) described a young yellow warbler with numerous pockets containing larvae: "Two of the lesions entered the pleural cavity, and the lungs had been partly devoured," which he considered evidence that when the larvae were numerous enough "they may literally devour the young birds alive." Rausch (1972) detailed the damage done to a fledgling pileolated [now Wilson’s] warbler by Protocalliphora larvae in subcutaneous cavities: "Soft tissues on both sides of the beak, including the right nostril, had been destroyed, and the resultant two dorsal grooves were continuous with subcutaneous cavities extending below the eyes bilaterally, .... The bilateral cavities, each occupied by a single larva, extended posteroventrad below the eye to the level of the external opening of the ear." The bird, perched motionless, was easily captured, refused food, and soon died, after which dissection revealed the damage. Bedard and McNeil (1979) recorded that a nestling savannah sparrow had lost an eye from larval action. The many records listed under P. braueri (q.v.) show the subcutaneous sites of the larvae, variously described as in boils, swellings, cysts, cavities, purulent sores, or lesions, and obviously some locations would have resulted in severe damage or even death. Shannon and Dobroscky (1924) listed a fragmented adult fly from Ontario, misidentified then as P. splendida aenea but now recognized as braueri, labeled as "from brain of a living fledgling of sparrow kind." Some years ago, the senior author identified an adult braueri reared by Murray L. Johnson of Tacoma, Wash., from a juvenile barn swallow, and Dr. Johnson (personal communication) kindly supplied the following information: "It was obviously very ill and unable to fly, though the feathers were well developed. There was a large cavity extending cranially from the right naris, containing two large larvae. The upper beak was weakened and movable; the right eye was closed." The bird died about a day later. Many labels and published information state that the larvae were on the head, and larvae in such a location could easily have done serious damage.

Host Relations and Ecology 31

In Europe, authors have also noted that infestation by braueri, especially if heavy, may kill young birds. Rognes (1985), in his recent revision of Trypocalliphora braueri, reviewed the European publications and records and concluded that "infestation weakens the host and is frequently lethal."

In conclusion, we believe that Protocalliphora \arvae do not ordinarily kill nestlings or fledglings, but under certain conditions they may do so, or they may so weaken the young that other factors will be lethal or will combine with larval feeding to cause death.

Host Relations and Ecology

All species of Protocalliphora are obligatory bloodsucking parasites of nestling birds, and they need more than one blood meal for maturation. Field studies and abundant available records suggest that, with the possible exception of species with wet and messy nests, all species of birds with altricial young (confined to a nest for some period of time) in the range of Protocalliphora will eventually be recorded as hosts of one or more species of Protocalliphora. If such birds within the range of Protocalliphora are not now known to be attacked, the dearth of records may result from (1) lack of collecting of the nests of those species, (2) low percentages of parasitism and reduced chances of finding infested nests, or (3) the nests may be absent Or rare in certain ecological niches. Many species of birds recorded from North America occur in geographical areas (e.g., Florida, Texas) outside the known range of Protocalliphora. On the other hand, precocial young birds would not be expected to be parasitized by Protocalliphora larvae, which need the relative permanence and stable food source of nestlings confined to a nest. Of course, even birds with precocial young could be hosts of Protocalliphora because of direct oviposition by P. braueri (e.g., there are records from domestic chickens and a budgerigar). Probably precocial young are not, or not often, as closely examined as nestlings.

The absence or virtual absence of host specificity, and the demonstrated occurrence of some species of Protocalliphora in the nests of a number of species of birds, make a list of hosts of less significance than formerly. For individual species of Protocalliphora, however, the host records could gradually

32 Protocalliphora

contribute to a typical or characteristic ecological picture for each species of fly, or of host preference if not host specificity.

Hall (1948) listed 51 species of birds reported as natural hosts of Protocalliphora in North America. To this list may be added 31 from later published literature and 57 (including three subspecies) formally published here for the first time (including dissertation records by Bennett and Whitworth), making a total host list at the present time of 139 for North America. In addition to these natural hosts, there are published records of Protocalliphora on two domesticated birds, the domestic chicken and the budgerigar, and from a nest of caged ringed turtle-doves used in experimental studies in Colorado. A list of the common and scientific names of the natural hosts will be found in the Appendix.

The ordinal distribution of the known hosts, shown in Table III, reveals the great preponderance of parasitism by Protocalliphora on the perching birds or Passeriformes. In terms of the percentage of species presently known to be parasitized by Protocalliphora, the passerine birds and the raptorial birds (Falconiformes and Strigiformes) are almost equally good hosts, with 41% for the Passeriformes (114 species out of 280) compared with 37% for the raptors (20 out of 54 species breeding in our region). Precociousness of young virtually precludes parasitism by Protocalliphora, hence the water and shore birds and the Galliformes are free of such parasitism, except for two unverified records of Protocalliphora (probably braueri) larvae on domestic chicks (Hearle 1938, Munro 1949). However, the young of water and shore birds are not usually handled and observed as Closely as those of nesting birds with altricial young, and if P. braueri deposits eggs directly on young birds as has been suggested (Rausch 1972, as P. hirudo), parasitism of the young of water and shore birds might occur in nature.

Table IV shows the distribution of records of Protocalliphora infestation among the families of perching birds, and it reveals the widespread occurrence of the parasitism and the sampling that has been done, either actively undertaken or achieved by the accumulation of scattered and chance rearings. If we disregard the skewed percentages that result from calculations involving only one to four species, the Table shows a range of between 17% and 47% for most families, very high for Hirundinidae (87.5%). The figures undoubtedly reflect the fact that most families of birds have some common species with readily observable or available nests that are easily sampled. Uncommon or secretive

Host Relations and Ecology 33

or inaccessible species are rarely sampled, and rare samples can easily be negative, especially if the percentage of parasitism is low for that species of bird or in that area or region. Swallows are common and readily observed, and hence most of our species have been sampled for Protocalliphora. The type and site of nest are factors: tight or solid cup nests or nests in cavities retain larvae and puparia of Protocalliphora more readily than "loose- weave" or few-stick nests, which may not provide suitable resting sites for larvae between blood meals, or from which larvae may drop to the ground and be missed by collectors or observers.

Mexico has the potential of a number of additional host species, but since no rearing records are known from that country Table IV is limited to hosts nesting north of Mexico, either as residents or as nesting migrants. In reality the number of available hosts should be lower and the percentages higher, because a number of chiefly Neotropical or subtropical birds occur along the Gulf Coast or in southern Florida and southern Texas, areas that are outside of what appears to be the normal range of Protocalliphora (cf. Map 1).

Bennett’s ecological study (Bennett 1957, Sabrosky and Bennett 1958) was the first broadly based investigation of habitat preference by species of Protocalliphora. His studies indicate that most species of the genus show habitat or ecological preference rather than host preference or host specificity. Different species of Protocalliphora tend to be found in different types of environment, and even in different levels or strata within those environmental types. Whitworth (1971, 1976) also studied the nest ecology of Protocalliphora but in such a diverse area that the data did not show habitat preference as much as nest site preference.

Authors differ on whether some situations should be referred to as host preference, habitat preference, or nest site (niche?) preference. Several species of Protocalliphora that have been extensively studied in the field (e.g., avium, asiovora, metallica, sialia) have a number of hosts but the hosts usually nest in the same type of habitat, e.g., in the canopy, near the ground, or in cavities. Probably it is technically a misnomer to speak of habitat preference; the birds prefer the habitat, the flies prefer the birds. Or do the flies also show preference for nest sites? Perhaps we may be pardoned if we continue to speak of habitat preference to avoid any circumlocution. For P. halli four hosts are known but barn swallow predominates. Whitworth (1976) reported that halli (as species IV) commonly infested barn swallow nests

34 —_ Protocalliphora

Table IIL Ordinal distribution of known natural hosts of Protocalliphora in North America north of Mexico

Orders of birds Total Sop.* Known as Hosts % Known Water and shore birds 211 0 0 Galliformes (grouse, quail, etc.) 21 oF 0 Raptorial birds: Falconiformes (hawks, 54 20 37 etc.), Strigiformes (owls) Columbiformes (pigeons, doves)** 1] 1 9 Cuculiformes (cuckoos, etc.) 6 ] 17 Miscellaneous orders: Psittaciformes 30 Ona 0

(parrots), Caprimulgiformes (nighthawks, etc.), Apodiformes (swifts, hummingbirds), Trogoniformes (trogons), Coraciiformes (kingfishers,

etc.) Piciformes (woodpeckers, etc.)*** 22 3 14 Passeriformes (perching birds: 280 114 4) blackbirds, buntings, sparrows, warblers, etc.)*** Totals 635*** 139"** 22

* Based on the AOU "Check-list of North American birds", 6th edition, plus Greenland species. The numbers include the species listed as "breeding" or "resident," including species non-native but established and therefore possible hosts of Protocalliphora. Extinct species are not included.

** Records from budgerigar, baby chicks, and ringed turtle-dove are not counted here, because they are unnatural hosts (captive audience!).

*** The figures include three recently classified as non-typical subspecies red-shafted flicker in Piciformes, Oregon junco and Audubon’s warbler in Passeriformes (Cf. Appendix).

Host Relations and Ecology 35

Table IV. Distribution of known hosts of Protocalliphora among the families of perching birds (Passeriformes)

Families Total Sop.* Known as Hosts %Known Aegithalidae (bushtits) 1 1 100 Alaudidae (larks) 1 1 100 Bombycillidae (waxwings) 2 1 50 Certhiidae (creepers) 1 1 100 Cinclidae (dippers) 1 1 100 Corvidae (crows, jays, magpies, ravens) 15 6 40 Emberizidae (buntings, blackbirds, 130 47 36 juncos, sparrows, towhees, warblers, etc. Fringillidae (finches, siskins, etc.) 15 7/ 47 Hirundinidae (martins, swallows) 8 7 87.5 Laniidae (shrikes) 2D 1 50 Mimidae (catbirds, mockingbirds, 9 3 33 thrashers) Motacillidae (pipits) 6 1 17 Muscicapidae (bluebirds, robins, 19 8 42 solitaires, thrushes, all in sub- family Turdinae, formerly Turdidae) Paridae (chickadees) 10 4 40 Passeridae (house sparrow) 2 1 50 Sittidae (nuthatches) 4 4 100 Sturnidae (starlings) 1 ] 100 Troglodytidae (wrens) 9 4 44 Tyrannidae (flycatchers, kingbirds, 33 13 39 pewees, phoebes) Vireonidae (vireos) 11 3 27 Totals 280 115 4]

* Tabulated as in Table IIL

36 Protocalliphora

bridges but did not infest cliff swallow nests in the same habitat (or nest site). P. parorum is a similar case, in that although it is known from nine different hosts, it is overwhelmingly a parasite of chickadees (Parus spp.), if we may lump three species of one genus as a kind of host preference. At one time, host specificity also seemed to be true for chrysorrhoea on bank swallows, although that might also have been interpreted as habitat preference (in a narrow sense) or nest site preference. Now that four other species of Protocalliphora are known from bank swallows, host specificity does not appear so reasonable. One might argue for either host preference (by selective oviposition) or nest site preference, and either would seem equally plausible, there being (with rare exceptions) only one species of bird in that one nest site or niche. Areas such as marshes, forest canopy, etc., are habitats in the broad sense of a certain type of environment, and they might also be nest sites in a narrow sense, although we prefer to think of a nest site as a particular place, or type of place, or niche, within a habitat.

Some species of Protocalliphora are known to parasitize many species of birds, but others are restricted to one or a few species of birds. The highest totals to date are by braueri on 42 species of birds (plus domestic chicks and a budgerigar), metallica on 35, and sialia on 33. These are common species often encountered in rearings, and all are parasites chiefly of small and ground- or low-nesting birds where the numbers of species of such birds as sparrows, warblers, and the Tyrannidae (flycatchers, kingbirds, phoebes) make possible the large totals. In contrast, avium is known from only 12 species because it is a parasite of large birds, chiefly raptors and high-nesting birds such as crows, with young confined to a nest for a long period of time, long enough for the relatively slow-growing avium to mature.

On the reverse side, some birds are parasitized by only one or two species of Protocalliphora, although some of this probably reflects lack of collecting. Again in contrast, three common birds with easily accessible nests that are most frequently sampled show 10 species on barn swallows and 9 each on American robins and common grackles.

Mixed infestations occur occasionally, and these can cause trouble in the correct association of sexes and of larvae or puparia. Bennett found 12.5% of 669 nests and Whitworth (1976) found 7.1% of 869 nests contained mixed infestations.

The host relations of braueri are especially interesting, as the species is often recorded in "tumors" or swellings beneath the skin, not only in nestlings and sometimes in fledglings, but at

Parasites of Protocalliphora 37

least once in a juvenile bird. It has been found in a wide range of hosts, from house finch to golden eagle. Presumably it may occur on almost any species of young bird. For further details, see the section on the adults of braueri. A similar special habit (see discussion under the species) is the infestation of ears and nostrils by avium and its western relative, asiovora, and sometimes by chrysorrhoea and sialia.

Parasites of Protocalliphora

Parasites and predators can be summarized briefly here. The most common is the polyphagous pupal parasite Nasonia [or Mormoniella] vitripennis (Walker) (Pteromalidae in Hymenoptera Chalcidoidea), which sometimes kills high percentages of Protocalliphora, as it does of many other muscoid flies. Morodora armata Gahan, another pteromalid, has been reared only from Protocalliphora, but it is seldom recorded. A third pteromalid, Dibrachys cavus Walker, was recorded as reared from Protocalliphora sp. by Burks (1979), but we have been unable to locate a North American record; possibly the entry was based on a published record in Europe. Finally, the encyrtid Tachinaephagus zealandicus Ashmead (Chalcidoidea) was recently recorded from Protocalliphora n.sp. by Gold and Dahlsten (1981).

Two other chalcidoid Hymenoptera have been reared from nests containing Protocalliphora but without definite association with it. The pteromalid Muscidifurax raptor Girault and Sanders was reared from a nest containing Protocalliphora asiovora, and three specimens of a eulophid, Pediobius sp. possibly alcaeus (Walker), were reared at Portage, Alaska, in 1978 (G. E. Haas) along with Protocalliphora hesperia from a bird nest, probably that of the gray jay, Perisoreus canadensis (L.). From the known hosts of the chalcidoids, it seems likely that they emerged from some other inhabitants of the nest, not Protocalliphora.

Evolution and Phylogeny

In the present state of knowledge of the group, with rapidly growing information about species in both Nearctic and Palearctic Regions, speculation on phylogeny seems premature. Attention has been concentrated on the difficult and presently more important problem of better understanding of the species

38 Protocalliphora

involved, with more precise definition for purposes of identification and recording of data.

As for the origin of the genus, it seems likely that Protocalliphora arose from a carrion-feeding calliphorid or calliphorid-like ancestor, and became adapted to feeding on living hosts with the availability of young birds in soiled nests. As James (1969) has pointed out in discussing the origin of parasitism, there is a logical transition from scavenging to preying on other organisms in the same habitat to feeding on young birds in the soiled nests. Within the calliphorid tribe Phormiini, there are suggestions of these stages, culminating in the obligatory hematophagy of larval Protocalliphora. Even within this genus itself there are stages from simple intermittent external bloodsucking, to penetration of external openings such as the ears and nares, to existence in subcutaneous cysts, though still bloodsucking.

The wide range of hosts and the usual lack of host specificity preclude any consideration of parallel evolution of hosts and parasites, such as has been done, for example, in parasitic mites and Mallophaga. Maggots of Protocalliphora are - with perhaps rare exceptions - not permanent parasites, only intermittent feeders, but they are obligatory parasites, and they will apparently feed on any birds with nests and nestlings. The availability of such a "captive audience" is the determining factor. Feeding on the blood of young birds may have developed relatively recently, probably from saprophagous larvae in birds’ nests to occasional or partial hematophagy and finally to obligatory hematophagy, but with specific association with certain host species. The parasites have by now become completely adapted to getting their blood meals from nestling birds and require blood meals for development. However, even though evolution has progressed to the point that parasitism on birds is obligatory and blood meals absolutely necessary for the parasites, experimental evidence shows that Protocalliphora larvae will also complete development on mammalian blood, including that of man, if a mammalian host is made available to them. Perhaps this is an indication that this last evolutionary step of obligatory parasitism on birds is relatively recent and has not yet become so firmly fixed but that the larvae can still develop on other bloods. Probably the isolating mechanism here is whatever attracts the female Protocalliphora to a nest for Oviposition, whether the hatching of the eggs, or the oxidation products from the egg remains, or the odor of the new-born young.

Collecting, Rearing, and Preserving 39

The close similarity of most species of Protocalliphora, the apparently high degree of habitat preference, and the occurrence of mixed infestations in nests suggest the possibility that hybridization might have at times played some part in speciation in the group. Perhaps the phylogenetic picture is partly reticulate evolution, rather than a dichotomous branching. But this is only speculation at this time, and moreover experiments by Bennett, while not exhaustive, indicated that hybridization was unlikely.

Collecting, Rearing, and Preserving Protocalliphora

It is surprising that many papers on parasites of birds, and even directions for collecting the parasites, make no mention of Protocalliphora. Attention is given to lice, fleas, louse flies (Hippoboscidae), ticks, and mites, but Protocalliphora larvae, in spite of the fact that they are fairly common and sometimes abundant, seem to escape notice.

Adults of Protocalliphora are seldom collected with a net, and then never in numbers. The most practical method of securing adults involves rearing them from larvae and/or puparia found in birds’ nests. This method usually makes possible a good series of specimens, samples of the immature stages, and a positive identification of the host, especially if the nest is located while the adult birds are still available.

Anyone interested in taking bird nests for scientific purposes, or in the removal of nests and use of substitute nests for parasite control (see next section on Control), must obtain a permit from the appropriate United States agency, the Canadian Wildlife Service at Ottawa, or a regional office of the Fish and Wildlife Service of the U.S. Department of the Interior. State or provincial permits may also be required.

Under most conditions, a nest should not be taken before the young birds have left it, although for experimental studies, or under certain circumstances where parasite control was desired in regularly observed nest boxes, nests have been removed and substitute nests provided. From the standpoint of the collector of Protocalliphora, early removal of a nest is both unnecessary and undesirable. With most birds, the period in the nest is short enough that the larvae of Protocalliphora require the full period to reach maturity. Ideally, an active nest should be located, the host identified, and the nest marked or noted for observation and collection at a later time. The nest should be examined for immature Protocalliphora as soon as possible after the fledglings

40 Protocalliphora

have left, because some larvae may drop to the ground to pupate. This is particularly true of birds that build small, loosely woven nests; in large or compact nests, most or all larvae will pupate in the nesting material. The nest should be torn into small pieces and searched thoroughly for the immature stages. In cup nests such as those of robin and barn swallow, the puparia may even be cemented into the mud of the nest. With loosely constructed nests such as those of the mourning dove, or with nests on the ground such as those of the white-throated sparrow, the ground immediately under and around the nest should be searched to a depth of three or four inches.

The recorded data should always include the locality, name of collector, and name of host. Other useful and desirable data are date of nest collection, date of adult emergence, habitat in which the nest was found, and height of nest above the ground.

Larvae of Protocalliphora, unlike those of most calliphorids, do not survive in damp surroundings. They are best maintained in dry sawdust until they pupate. Large larvae about to do so are recognizable by the lack of contents in the alimentary tract, and by a uniform gray, cream, or yellow color. Other large larvae (10-15 mm) will pupate also in three to four days. A sample of the larvae should be preserved for study by placing them in a KAAD solution (1 part kerosene, 10 parts 95% ethyl alcohol, 2 parts acetic acid, 14 parts dioxane) for 24 hours, and storing in 70% ethyl alcohol + 10% glycerine. If KAAD solution is not available, the larvae should be killed in hot water before storing in preservative.

Puparia being held for emergence of the adults are best maintained in a mixture of sawdust and boric acid (about 1% of boric acid powder), slightly wetted. Ideally, some or all of the puparia should be segregated in separate vials, so that each emerging adult can be associated with its own puparium. Individual adults should be kept alive for 24-48 hours after emergence, for maturation and hardening of the integument before they are pinned. Adult and entire puparium should be kept together. The puparium and the two halves of the cap can conveniently be stored together in a gelatin capsule, which can then be pinned beneath the adult. Empty puparia should be Stored dry.

The accurate association of adult and immature stage is of great significance. It may be critical in the differential analysis of mixed infestations, and in the study of incompletely known species and species complexes, such as the western fauna at the present time. Here the taxonomist is at the mercy of the

Control of Protocalliphora 41

collector. If adults and puparia from a given nest rearing are heterogeneously associated, but only one species happens to be present, no confusion will result. On the other hand, improper association in mixed infestations, especially if only a few puparia are preserved, can lead to erroneous conclusions. If accurate association of puparia and eclosed adults is not possible or practicable, then puparia should be preserved separately so that the taxonomist will be aware of the lack of individual relationship. He can thus deal with a series of adults associated with a series of puparia. If both series are homogeneous and of reasonable length, he has a useful association upon which to base conclusions and further study, despite the lack of individual associations.

Control of Protocalliphora

Although our studies have not been concerned with control of blow fly infestations, we give here a review of suggestions that have appeared in the literature. Obviously, control in nature for such flies as Protocalliphora is impracticable in most cases, as well as unnecessary. Furthermore, migratory birds ordinarily would not and should not be disturbed in their nests and the need for control would not be apparent. Where nest boxes are maintained and cleaning operations show that the birds were heavily attacked, some might wish to try to control the blow flies in order to alleviate the worst of the attacks and to promote survival of the young birds in the next brood or the following season.

Regular removal and destruction of nests immediately after the young birds have fledged would destroy larvae and pupae of Protocalliphora and might tend to reduce the local population and perhaps hold it down to reasonable levels, although we question any significant or lasting effect on the total area population of the flies. Unfortunately the procedure would also destroy the chalcidoid parasites, which are very common and often kill 25-100 percent of the puparia in a nest. Nest destruction would seem to have the greatest impact on hymenopterous parasites as the season advances, when the percentage of parasitism usually increases. The commonest parasite, Nasonia vitripennis, is common everywhere on a wide variety of muscoid hosts, and probably removal of a comparatively few would have only a slight local effect and no serious or lasting inhibition on the area population of the chalcidoids.

42 Protocalliphora

Another method of combating parasitism by Protocalliphora larvae was reported by Johnson (1932), based on information from one of his correspondents. The nest of the first brood of birds was removed and a handmade nest was substituted. This removed any larvae as well as any fly eggs that had just been laid, and gave the young birds a clean start. Such a method, however effective, would be time-consuming and could only be used in a limited way on nests or nest boxes that were being closely observed or being used in experiments or life history studies. It would also not deal with eggs laid directly on the nestlings.

Zeleny (personal communication) recounted to the senior author his experiences with nest removal, which were the basis of his recommendations in Zeleny (1970). In a closely observed nest, young bluebirds 10-12 days old were weak, and something seemed to be amiss. Inspection showed the nest to contain over 100 maggots of Protocalliphora, and it was therefore removed and a clean nest of dry grass was substituted. The young perked up and fledged successfully. Another infested nest was likewise successfully treated.

Some authors have recommended dusting the birds’ eggs with a mild insecticide, a mild flea powder (Highhouse 1963), pyrethrum powder (Brower 1966), or rotenone dust (Zeleny 1970, 1976), in order to control the fly larvae before the chalcidoid parasites are present. The timing would depend partly on the duration of effectiveness of the insecticide. For example, pyrethrum dust has a residual life of about one hour, perhaps two to four hours in a dark nest box. If fly eggs are not laid until after the birds’ eggs hatch (cf. life history studies), there would be no fly larvae present to be killed if the nests were dusted too early; if fly larvae were present, dusting would have to be very thorough because only larvae actually contacted with the insecticide would be likely to die. Note the "tough" larvae mentioned in the section of life history studies.

Zeleny (1976) has given a well considered discussion of the possible use of insecticides, listing 1% rotenone powder, pyrethrum powder, and pyrethrins. Pyrethrin formulations especially for use on caged birds could be used in nest boxes. All writers caution that although the insecticides mentioned do not appear to harm the birds, they might have obscure or deleterious effects that are not presently known, and hence they should be used sparingly and not routinely. Zeleny stated that "the dusting may be done before or after the eggs have been laid, but before they have hatched," although no fly larvae will be present then

Control of Protocalliphora 43

and any effectiveness will depend on the duration of the insecticide’s potency. He does offer a further thought: "If it is necessary to treat a nest after the young birds have hatched, the nestlings should be lifted gently from the nest before the application, and the insecticide should be worked carefully into the nesting material with the finger so that as little of it as possible will come into contact with the nestlings when they are put back." No doubt this would be effective, because the larvae leave the nestlings after feeding and withdraw into the nest material where they could come into contact with the insecticide. We believe that under most conditions and for most people, however, birds in nest boxes should be left undisturbed and unexamined for parasites. If examination of the nests after the young have fledged showed serious blow fly infestations, then the nests could be destroyed and the boxes cleaned out, and perhaps the following season or for the following brood dusting could be done in the manner suggested by Zeleny.

A reader desiring to control Protocalliphora larvae in nest boxes or under experimental conditions would do well to check with a veterinarian specializing in or at least concerned with bird care. Sevin (carbaryl) is commonly used by veterinarians to control parasites on adult domestic birds, with no ill effects, and perhaps this would be effective in a nest box although we do not know that this has been tested. This product has about a one-week residual effect, so its effectiveness would be longer lasting than pyrethrum, for example, if it proved to have no ill effects.

Two factors need to be kept in mind. First, infestation by blow fly larvae may not be detected without undesirable disturbance to the birds. Second, larvae of P. braueri may stay attached to the host and would not be removed with the nest, if that method were used. Larvae that may stay in the nares or ears, such as those of Protocalliphora avium and asiovora, would also remain on the birds, of course, but birds so infested (raptors, corvids, bank swallows, starlings) are not those involved in nest boxes.

A side effect of all the methods suggested would be the reduction or even elimination of mites, fleas, and lice, no doubt to the delight and better health of the nestling birds.

44 Protocalliphora

Taxonomic Section Genus Protocalliphora Hough

Protocalliphora Hough, 1899a: 65-66. Two species. Type species, Musca azurea Fallén, by original designation.

Avihos pita Hendel, 1901: 29. Five species. Type species, Musca azurea Fallén, by original designation.

Protocalliphora (syn., Avihospita); Aldrich, 1901: 68; Aldrich, 1905: 523.

Apaulina Hall, 1948: 179. Ten species. Type species, Protocalliphora avium Shannon and Dobroscky, by original designation.

Protocalliphora (syn., Apaulina, ex Sabrosky in litt.); Hamerstrom and Hamerstrom, 1954: 5; James, 1955: 24; Zumpt, 1956: 94.

Protocalliphora, subgenus Orneocalliphora Peus, 1960: 198. Four species. Type species, Musca chrysorrhoea Meigen, by original designation.

Trypocalliphora Peus, 1960: 199. Three species. Type species, Avihospita braueri Hendel, by original designation [Sabrosky 1967: 122 was curiously and inexplicably in error in citing T. lindneri Peus as the type species].

Protocalliphora (syns., Apaulina, Orneocalliphora, Trypocalliphora); Zumpt, 1965: 83.

Trypocalliphora; Rognes, 1985: 371-382. [Revision].

Protocalliphora; Shewell, in McAlpine et al, 1987: 1134-5, 1140, 1143. The generic diagnoses of Hall (1948, as Apaulina) and Zumpt

(1956) will serve except for the few characters in which Apaulina

was said to differ from Protocalliphora. Briefly, Protocalliphora

can be characterized as follows in the family Calliphoridae: Predominantly metallic blue to bluish purple or bluish black, with a few species partly aeneous or cupreous, all somewhat gray-microtomentose; epistoma strongly warped forward but not elongate, ending only barely below level of vibrissae; scutum in most species more or less flattened centrally, and with long, strong, erect bristles, including several pairs of well developed presutural acrosticals; greater ampulla (subalar know) with naplike pile, not long haired; stem vein (remigium, basal section of R) ciliate posteriorly on upper surface (Fig. 4a); subcostal sclerite setulose (Fig. 4b); surface of calypteres with short to minute pile but without long hairs, the lower calypter bare and narrowly rounded; hind coxa bare posteriorly. The side view of the head and the wing venation, which are quite uniform for the genus, have been figured by Hall (1948, plates 4C and 9H).

Taxonomic Section 45

The flies are readily recognized as typical calliphorids (blow flies), the color (except in a few species) being suggestive of Calliphora but the body shape narrower as in Phormia regina (Meigen). Detailed species descriptions of color are usually difficult and unsatisfactory because of the range of condition of the specimens. By far the largest part of the available material has been reared, and the specimens were killed in different degrees of maturity and condition. The thorax and abdomen are dark metallic bluish or purplish black in males, the females likewise in some species but green in others (Figs. 33-35). Males are always more shining than the females, the latter being more heavily microtomentose, at least on the scutum, with the microtomentum as viewed from behind demarcating three broad but weakly defined shining mesonotal stripes. These stripes are especially evident when the thorax is viewed from behind at a low angle. The head is chiefly dark, but the frontal vitta and genal grooves are usually lighter, reddish to dark reddish or brown. The lower ends of the facial ridges and the vibrissal angles are usually reddish yellow, and the palpi, narrow bases of 3rd antennal segments, and the 2nd antennal segments at least apically, are reddish to reddish yellow in most species.

The general color and structure are surprisingly uniform throughout the genus, a uniformity that has led to numerous misidentifications. Hall (1948) described in great detail the characters held in common by all calliphorids. His characterization of adult Protocalliphora (as Apaulina in his work) will serve for all species of Protocalliphora herein treated, except for a few characters, discussed elsewhere, in which we do not agree that Apaulina is distinct from Protocalliphora. The general characterization has not been repeated, and the keys and descriptions in this revision are focused on features that have shown useful differences. For ready reference by users of this work, Figures 1-4 illustrate the major structures and measures that are used.

One character, the strong mid ventral bristle on the middle tibia, is present in all known Nearctic species and has not been mentioned in the descriptions. Protocalliphora maruyamensis Kano and Shinonaga (1966), described from Japan, is unique among all known species in lacking this bristle. Also, females of Protocalliphora regularly have one pair of reclinate upper orbital bristles (Fig. 1), and the character is mentioned only for braueri, in which the bristles are absent.

The wing venation is uniform throughout the genus and is not referred to in the descriptions. Hall (1948) has described it, and

46 —_- Protocalliphora

figured the wing of P. metallica (his plate 9H). We note only that the 3rd vein (R 5) is actually setose from a third to halfway to the small crossvelit (r-m), not merely at base.

An occasional author has synonymized Protocalliphora under the older name Philornis Meinert 1890, probably because both attack nestling birds. The latter is a genus of Muscidae quite different from the calliphorid genus Protocalliphora, and references to those authors have not been included here. Philornis is chiefly Neotropical, with outliers in Florida and southern Texas.

Synonymy of Apaulina

Apaulina was proposed on the basis of supposedly significant characters of chaetotaxy, reinforced by zoogeography, so that it is important to examine the basis for our synonymy, especially since Protocalliphora was then said not to occur in North America. Additional material has revealed that the characters used for Apaulina are actually somewhat variable, that there are intermediate species, and that a typical species of Protocalliphora even in Hall’s restricted sense does in fact occur in North America, in the nests of a Holarctic bird, the bank swallow. The points of supposed differentiation are discussed seriatim.

1. Two posterior bristles on the fore tibia in Protocalliphora, usually one in Apaulina: Occasional specimens of Protocalliphora have only one, and conversely, certain species of Apaulina (especially hesperia, sapphira and tundrae) quite often have two, at least on one side. Occasionally two have been seen even in avium and asiovora.

In view of the considerable reliance placed on the character of the posterior bristles on the fore tibia, the variability in an obviously homogeneous series of P. tundrae reared from snow bunting nests on Ellesmere Island is significant and enlightening. The series consists of 70 specimens (32 males, 38 females), of which 36 (19 males, 17 females) have two posterior bristles on each fore tibia, and 22 others (2 males, 20 females) have two on one side or the other, but 12 (1 male, 11 females) have only one on each fore tibia.

2. Tympanic membrane (Fig. 3b) haired or setulose in Protocalliphora, bare in Apaulina: Typical species of the former, such as azurea and chrysorrhoea, do have a number of strong black setulae, and Apaulina spp. usually none, but some species included in the latter regularly have some hairs on the tympanic membrane. For example, out of 71 specimens of Nearctic braueri

Taxonomic Section 47

checked for this character, 28 (39 %) showed from one to 13 hairs on one or both sides. One individual from Seattle, Wash., had 13 on the left side and 10 on the right, all coarse, black, and conspicuous. Even in such a distinctly Apaulina-like species as metallica, occasional individuals have up to four or five tympanic hairs. A reverse combination occurs in the Nearctic species /ata, which has the two posterior bristles on the fore tibia, typical of Palearctic Protocalliphora, but only a few (2 or 3) weak hairs on the tympanic membrane.

3. Postalar wall (Fig. 3b) with a median tuft of hairs in Protocalliphora, bare in Apaulina: Hall (1948) regarded this as an important character for Apaulina, and Peus (1960) considered that the question of whether Protocalliphora can be used for Nearctic species depended on the taxonomic value assigned to this very character. Both believed, however, erroneously as we now know, that the postalar wall is bare in all the Nearctic species. Many if not most specimens of Apaulina actually have one to a few hairs on the wall, although these admittedly are usually short, pale, and inconspicuous. In some individuals part of these are dark, and occasionally long. Moreover, P. chrysorrhoea, a widely distributed species in Europe with a conspicuous tuft of hairs on the postalar wall, is now known to be Holarctic, a fact unknown to both Hall and Peus. This further weakens the argument that Nearctic and Palearctic species can be neatly divided by this character.

4. Scutellum with apical bristles (Fig. 3a) in Protocalliphora, without apicals in Apaulina: Apparently the same pair of bristles is present in almost all specimens, but varying in position from nearly apical to subapical, and sometimes even appearing to be discal scutellars.

5. Stem vein (basal section of R, remigium) with some cilia on ventral surface in Protocalliphora, none in A paulina: Generally true, but sapphira and chrysorrhoea in North America often have the setae, and braueri in Europe does not, so there are exceptions on both sides. The cilia have also been seen on occasional specimens of other Nearctic species.

6. Immature stages: There are no generic differences in the immature stages between the species of Protocalliphora and Apaulina Hall (1948). Comparison of species results in even closer ties. For example, the puparia of chrysorrhoea are similar to those of sialia, but adults of the latter are typical Apaulina sensu Hall and those of the former Protocalliphora in the strict sense.

7. Zoogeography: The difference once thought to exist

48 Protocalliphora

--Protocalliphora Palearctic and Apaulina Nearctic--has disappeared as more material has become available. Two species have been found to be Holarctic: braueri, formerly called hirudo in North America and lindneri in Europe, and chrysorrhoea, which occurs in bank swallow nests in both Palearctic and Nearctic Regions. (In fairness, both chrysorrhoea in North America and the hirudo-like species in Europe were unknown to Hall when he proposed Apaulina). Hall’s new species sapphira was described in Apaulina, but the tympanic pit has coarse black setae and the holotype and one paratype have two posterior bristles on the fore tibia. The long series of tundrae now available shows that these characters are typical of many--but not all--examples of the species. P. /ata from North America has virtually bare tympanic pit but two posterior bristles on the fore tibia, hence a combination of the characters supposedly characteristic of Apaulina and Protocalliphora, respectively. The reverse combination is found in P. asiatica Zumpt in the Palearctic Region, which has numerous black setae on the tympanic pit but only one posterior bristle on the fore tibia. It is known from only a single specimen, and the tibial chaetotaxy could be variable, as often noted in the Nearctic hesperia, for example.

In conclusion, then, the variation or intergradation of the characters of chaetotaxy, and the absence of the supposed geographical separation, together with the fundamental similarity of the anatomy, male terminalia, immature stages, ecology, and host relationships of the Palearctic and Nearctic species ali lead us inevitably to the conclusion that Apaulina is synonymous with Protocalliphora. It cannot even be recognized as a subgenus.

Synonymy of Orneocalliphora

Peus (1960) proposed to divide the genus Protocalliphora into two subgenera: Orneocalliphora for those species (e.g., chrysorrhoea) in which the two sexes are essentially alike in size, color, and microtomentum, and Protocalliphora in the strict sense for those species (azurea et al.) with strong sexual dimorphism in size and microtomentum, and usually in color. In the numerous Nearctic species there are different degrees and combinations of characters, and we do not believe this subgeneric division has any validity. Zumpt (1965) and others have also rejected the division as untenable.

Taxonomic Section 49 Status of Trypocalliphora

Peus (1960) also proposed a new genus 7rypocalliphora for three presumably distinct species, the Palearctic lindneri and braueri and the Nearctic hirudo. As with Apaulina and Orneocalliphora, our review of the numerous Nearctic species of bird blow flies leads us to question the taxon as a genus. A number of the characters used are differences only of degree, of the amount of reduction from typical Protocalliphora. For example, the range in amount of flattening of the scutum is so great in some series that one is led to suspect the influence of degree of maturity, especially since so many available specimens are reared, and because the relatively few wild-caught late-season specimens, obviously mature, are quite convex. This is especially true in the series of metallica, the species most often net collected late in the season. The postalar wall is said to be bare in Trypocalliphora, and it usually is, but specimens have been seen with one to three short black hairs, including several specimens from Schlorer’s original rearing of Jindneri at Diersheim, Germany. Furthermore, in ordinary Protocalliphora there is a range from a strong and distinct tuft of rather long hairs to few or none, and Apaulina was said to be characterized by having the postalar wall bare. The tympanic tuft varies through the same range (cf. previous and also later discussion of that character). The surstyli of the male terminalia range from extremely narrow, long and curved, to the broadly obovate type of several American species (cf. Figs. 7 - 32). The frontal width in the male likewise ranges from very narrow to broad (Figs. 33-41), in different combinations with the male terminalia. Species may or may not be dichromatic. Puparia, and hence the mature larvae, are all of the same general type (Figs. 42-64), but the integument ranges from smooth and almost without spines--the spines being reduced to minute tubercles as in metallica--to densely long spined, with different degrees of rugosity and cuticular folds. Some larval characters thought by some European authors to be characteristic of Trypocalliphora appear to us to be only degrees of reduction from Protocalliphora, extreme though they are.

Grunin (1970a) in his generic key characterized the calypteres of Trypocalliphora as yellow with yellow edge but those of Protocalliphora as white with light edge, at most the thoracic calypter brown. That was probably a character of convenience for the species with which he was dealing rather than a significant generic character. There is again a range in the North

50. ~—- Protocalliphora

American species that casts doubt on the generic value of the character. In Jata and brunneisquama, the calypteres are very dark, black or blackish brown. In cuprina and hesperia they are brown. In aenea and halli, they are white or at most yellowish white in the females but deep yellow (aenea) to light brown (halli) in the males. In braueri the calypteres of the males are darker than those of the females, a deep yellow to light brown, compared with yellow for the females.

Some European authors have maintained that Protocalliphora and Trypocalliphora are distinct because species of the former are external bloodsucking parasites whereas those of the latter are subcutaneous; however, the subcutaneous habit is neither uniformly nor exclusively characteristic of the species referred to Trypocalliphora. It is important first to note that the species of Trypocalliphora are also bloodsucking, even though they commonly embed themselves when they feed. In Nearctic braueri, atypical Trypocalliphora, Bennett has found many larvae in the nest material, engorged with blood, like typical species of Protocalliphora. It is true that records of subcutaneous larvae in North America frequently prove to be braueri, but not always. P. avium and P. asiovora are often found in the ears of raptorial birds and may even penetrate to the brain, but they are also found embedded in the flesh in other areas (cf. notes under avium in particular; also Hamerstrom and Hamerstrom 1954). Specimens of the new species seminuda, atypical Protocalliphora, were reared along with braueri from larvae found in purulent sores on a fledgling horned lark in New Mexico (Walton 1914). Specimens of chrysorrhoea were reared in Alberta "from larvae that were removed from under the skin and breast muscles of nestlings."

Generic distinction between Trypocalliphora and Protocalliphora breaks down notably in P. deceptor, which is intermediate. The habitus of this species is immediately suggestive of braueri. It is strongly dimorphic in the color of males and females. The scutum posteriorly is convex. The frons is unusually narrow in both sexes, little over half the width of an eye in the female and so narrow in the male that the parafrontals almost touch. The basicosta is yellow to bright orange. Unlike Trypocalliphora, however, the reclinate upper orbital bristle ("Frontalborste" of Peus) is present in the females. This alone might not be considered a _ serious divergence from Trypocalliphora, but the male terminalia are of the type found in avium and asiovora, with broadly obovate surstyli (Fig. 9) and aedeagus typical of subgenus Protocalliphora (Fig. 6a).

Taxonomic Section 51

Furthermore the puparium is covered with numerous fine spines, quite unlike the relatively bare puparium of braueri.

Protocalliphora aenea likewise shows, at least in mature specimens, a convex scutum, and it has a strong sexual dimorphism in color, but the reclinate upper orbital bristle is present in females, the front is wider in both sexes, the male terminalia are more like those of chrysorrhoea, and the puparium is well covered with spines.

In conclusion, the various combinations of characters lead us to doubt that more than one genus can be recognized. We do agree that the species described in or referred to Trypocalliphora-- originally three, lindneri, braueri, and hirudo, now one, braueri, with the others as synonyms--form a species group characterized notably by the distinctive aedeagus and in the female by the lack of a reclinate upper orbital bristle. In our view, variation and the range of characteristics in the now greater known number of species greatly decrease the apparent distinctiveness of Trypocalliphora although it can be argued that the distinct aedeagus is still justification for its recognition. In order to preserve the usefulness and meaningfulness of the name Protocalliphora in general biological literature but at the same time to segregate the species group, we choose to recognize Trypocalliphora as a subgenus of Protocalliphora. Sabrosky (1967) has already suggested this. The remaining species form the typical subgenus Protocalliphora.

Rognes (1985) published a detailed study of Trypocalliphora (as a genus) in which he concluded that there is only one species, wide-ranging and Holarctic. The oldest available name is Hendel’s Avihospita braueri (1901), with synonyms Protocalliphora hirudo Shannon and Dobroscky 1924, T. lindneri Peus 1960, and 7. compacta Grunin 1966. This synonymy returned to the identification by Sabrosky (Heinz 1954) of hirudo for European specimens, which was accepted in _ several publications such as the revision by Zumpt (1956). (At that time, the specific identity of braueri was unknown). Peus (1960) rejected the identification of hirudo and he described European specimens as Jindneri n.sp., which was accepted by Zumpt (1965). Rognes (1985) has concluded that lindneri and hirudo are in reality conspecific, that /indneri was based on variable characters, and that both are synonyms of the older but poorly described braueri. We cannot contribute on the last named, but Rognes thoroughly studied the extant type series. We do concur in the conspecificity of European and North American material, and

52 Protocalliphora

we have adopted the name braueri in place of the familiar and rather often cited hirudo.

Generic Relationships of Protocalliphora

There are few clearcut generic distinctions in the adults of Protocalliphora, as one quickly realizes when trying to construct a satisfactory generic key. However, the characteristic biology, the obligatory bloodsucking parasitism on nestling birds, and the distinctive larvae mark the group as a distinct entity, although some might argue that it is merely a specialized subgenus of Phormia.

The larvae are more distinct generically than the adults. Larvae of Phormia resemble those of Cochliomyia and other chrysomyine calliphorids in having anterior bands of spines on the abdominal segments (screwworms). In Protocalliphora, on the contrary, the spines are distributed over most of the surface, even in a few species in which the spines are extremely reduced (metallica, braueri). Furthermore, all species of Protocalliphora have in all larval instars the characteristic and unique feature--occasionally much reduced but detectable--of a fringe of long hairs on the anterior margin of the prothoracic segment (Hall 1948, plate 38D). These larval features and the characteristic biology thus justify generic recognition, even though the adult characters may not seem impressive.

Protophormia and Boreellus, which contain species also formerly included in Phormia, are easily separated from Phormia and Protocalliphora, but the latter two are not so easily separated from each other. Hall (1948) keyed Phormia regina as having "Mesonotum convex; eyes in male very narrowly separated or contiguous; mesothoracic spiracle with bright orange hair," whereas the other phormiines, including all Protocalliphora (as Apaulina in his book), were said to have "“Mesonotum conspicuously flattened on disc; eyes in male distinctly separated (except in several species of Apaulina); mesothoracic spiracle with dark hair (dark orange brown in Apaulina_ hirudo)." Unfortunately, none of these characters will consistently differentiate Protocalliphora from Phormia. More species of Protocalliphora are now known in which the eyes of the male are only narrowly separated, and in which the mesothoracic spiracle has brightly colored hairs, not as bright as in Phormia regina but enough to be confusing. As for the first character, some difference of interpretation is evident. Hall, following Townsend (1935), keyed Protocalliphora as having the postscutum, or

Generic Relationships 53

posterior portion of the mesonotum, "conspicuously flattened on disc," but under this they were including P. hirudo, which was included by Peus (1960) in his genus Trypocalliphora, which he characterized as having the mesonotum convex. In practice, the character is difficult to apply because specimens are so often pinned through that part of the body, either in the center or to one side of the midline. Obviously also there is some difference in the degree of flattening. Further, when one compares reared material--all too often more or less teneral--with specimens caught in nature, one sees that the flattening of the mesonotum is often more obvious in reared material and scarcely or not at all evident in hand-caught specimens. Then when one remembers that most specimens in collections are reared, because these flies are seldom collected otherwise, one realizes that most specimens in collections will show flattening. There may be slight differences between species in the degree of convexity of the mesonotum, if one could get perfect specimens and measure the curvature properly, but we have concluded that much of the observed flattening is an artifact, or is more apparent than real, and that in practice it cannot be used to separate Trypocalliphora from Protocalliphora. This conclusion is reinforced when one compares braueri with deceptor: both with convex scutum but radically different male terminalia, and female with reclinate upper orbital bristle in deceptor.

Discussion of the status of the other genera of the tribe Phormiini is irrelevant here; it is sufficient to point out in the following statement the distinctive nature of Protocalliphora compared with related phormiines:

Protocalliphora. Upper (alar) calypter with at most minute pile on the upper surface, any long hairs confined to the rim; anterior margin of prothoracic segment of larvae with a complete band of spines, forming a characteristic fringe, which is long in most species (Hall 1948: plate 38D), greatly reduced in braueri; larva dorsally and laterally thickly set with spines over the entire surface, except in a few species in which they are greatly reduced; larvae obligatory bloodsucking parasites on nestling birds; dorsum of thorax and abdomen more or less microtomentose, the scutum viewed from behind (at least under a microscope) showing three dark stripes, more conspicuous in females than in males; mesonotal bristles strongly developed, including complete rows of acrostical bristles and 3 or 4 strong post-intraalar bristles (Fig. 3a); postalar wall often bare or with relatively few inconspicuous hairs about midway, especially in Nearctic species (Fig. 3b).

54 __ Protocalliphora

Phormia, Protophormia, and Boreellus: Upper (alar) calypter with numerous long, erect hairs on upper surface of the outer half or more (pale and easily overlooked in Phormia, black in the others); larvae without prothoracic fringe; most larval segments with distinct anterior or anterior and posterior bands of spines, the intervening integument bare; larvae normally saprophagous, occasionally involved facultatively in myiasis; thorax and abdomen in both sexes highly shining, only a thin, obscure microtomentum on the anterior slope of the scutum, the scutum not vittate; acrostical bristles absent (Protophormia, Boreellus ) or weakly developed (Phormia); two post-intraalar bristles, rarely (Protophormia) a weak third; middle to lower half of postalar wall with numerous, long, coarse, conspicuous hairs.

Possible confusion of Protocalliphora with the blue bottle flies, Calliphora or Cynomyopsis, because of the blue color of the abdomen, is easily resolved by a glance at the upper surface of the stem vein (basal section of R, remigium), which is entirely bare in the calliphorine genera but long ciliate posteriorly in the phormiine genera (Fig. 4a).

The place of Protocalliphora in the blow fly family Calliphoridae and its relationship to the genera previously discussed may best be shown by a key, somewhat abbreviated for present purposes. The characters used for the adults, although not appearing of great significance, will serve to separate Protocalliphora from Phormia. Superficially, Phormia is easily recognized by having a dark and much more shining and unstriped thorax, hence its common name, black blow fly. Species of Protocalliphora, even in the relatively shining males, have a more microtomentose thorax, the microtomentum especially evident in two broad submedian stripes that delineate three broad shining stripes, particularly when viewed from behind. Protocalliphora is also much more bristly, and the bristles are larger and more erect.

In passing, we note that Lehrer (1970) proposed a new tribe Protocalliphorini, but we agree with Shewell (1987) that this tribe is not justified. The fundamental similarity of Phormia, Protophormia, and Protocalliphora indicates to us that the last named is properly placed in the Phormiini.

Generic Relationships 55

Abbreviated keys to genera and higher categories of Nearctic Calliphoridae Adults

1. Stem vein (remigium) ciliate posteriorly on upper surface of wing (Fig. 4a) (Subfamily Chrysomyinae) ........ 2

-- Stem vein (remigium) bare above ..................- RNY ea eu el) Sey RR i Subfamilies Calliphorinae (blue- and greenbottle flies) and Polleniinae (cluster flies)

2. Hind coxa bare posteriorly; lower calypter at most with short, naplike pile above; head predominantly black (Tribe PORE) RAR 1 Sr ee Was eh owl ah ae gs MR 3

-- Hind coxa setose posteriorly; in Nearctic forms, lower calypter with long erect hairs above on basal half; head predominantly yellow......... Tribe Chrysomyini

3. Presutural acrostical bristles present, strong (Fig. 3a); head not elongate below, the vibrissal angles not strongly projecting and lower margin of head in profile appearing FOUMGSG! re Mr eaten ae tics We oe ean Gat MLA lS eel 4

-- Presutural acrosticals absent or vestigial, at most only faintly thicker than the long, fine, erect scutal hairs; head strongly elongate below, the vibrissal angles projecting and vibrissal axis obviously longer than antennal axis, lower margin of head straight ....... Protophormia

. Townsend and Boreellus Aldrich and Shannon

4. Three postsutural intraalar bristles (Fig. 3a); postalar wall (Fig. 3b) bare or with few minute hairs in most Nearctic species (long, but still fine and weak, in chrysorrhoea); dorsum of abdomen more or less heavily microtomentose; larvae are bloodsucking parasites of nestling birds.....

RN tee Rane ad Aha pa Hees vebre Protocalliphora Hough

-- Two postsutural intraalars, the first absent; postalar wall with strong tuft of long bristly hairs; dorsum of abdomen entirely shining; larvae normally saprophagous but may have healthy tissue in wound myiasis ............

IONS tar RN SAN Phormia Robineau-Desvoidy

56 Protocalliphora

Mature larvae and puparia

1. Prothoracic fringe (Hall 1948, pl. 38D) present (very short in braueri); integument dorsally and laterally densely covered with spines in most species, minute (like fine sandpaper) in metallica, minute and reduced to anterior spine band in braueri; posterior surface (Fig. 42ff) with numerous spines except in braueri and metallica

LARS ASE ae eS Protocalliphora Hough

-- No prothoracic fringe; most segments with anterior band of strong spines, a few may have anterior and posterior bands, rest of integument bare (screwworm appearance); posterior surface bare of spines . Other Calliphoridae

Taxonomy and Taxonomic Characters of the Adults

In the descriptions of the species, the following arrangement has been adopted: Citations and synonymy, brief diagnosis, description of male, description of female, description of immature stages, type series, specimens examined (figures for males and females), distribution (general statement, followed by records, which are detailed if only few, more generalized if many), hosts, ecology and biology, variation, remarks and etymology (for new species).

References under each species are chronological, except in one or two cases where minor references were grouped under different combinations, each group chronological within itself. For citations other than original references, the name of the

Taxonomy of Adults 57

author of the species is omitted; the name of the species is then followed by .--, and the author and citation of the reference.

In the descriptions of the adults, all measures were based on 25 specimens of each sex except where stated otherwise. Occasionally, damage to a specimen made some measure(s) unavailable, and for some new species the limited available material is admittedly an inadequate sample. All measurements were made by means of an ocular micrometer, at the same magnification (27x). The mean ratio between any two measures is based upon the sums of the measurements for each. Each mean ratio is followed by a statement in parentheses of the range of the individual ratios. When fewer than 25 specimens were measured, ranges are followed by a slant and the number of specimens measured. Thus for males of beameri, the ratio of frons: head width is 0.12 (.11-.14/10), based on 10 specimens. The same method is used when 25 specimens were measured but one ratio was based on a smaller number, e.g.,/24.

Under Distribution, after the general statement, localities are arranged by state and province in the order Alaska, Canada, Contiguous United States, and Mexico. Occasionally the records are separated into western and eastern records for purposes of special discussion. Identifications of adults are by Sabrosky and of larvae and puparia by Bennett unless stated otherwise.

For seasonal range, available data may be summarized, but the information is not considered significant and dates have usually not been given. The larval period corresponds to the nesting period of the bird hosts, and this governs the appearance of adult flies. However, the latter are long lived and apparently overwinter, and thus there are records of occurrence outside of the nesting season. Available records are also complicated by the frequent failure to specify on labels whether the dates given are those of collection of nests or of emergence of adult flies. Any unusual dates are noted.

Many factors make this genus of closely related species a particularly difficult one. There is some variation, often wide, in all the characters of chaetotaxy and proportions (e.g., note the ranges in Tables V and VI) that one must use because of the paucity of effective differences. Some of these characters are further modified by the degree of maturity of individual specimens. Measurements given in this paper are based on examples judged to be mature (i.e., fully extended and hardened). Body color is little affected by slight immaturity, and its correct interpretation is rarely jeopardized. If a specimen is greasy or dirty, or has become wet while in a killing jar or while being

Protocalliphora

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Taxonomy of Adults 59

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relaxed for extraction of the male terminalia, the color of the calypteres is generally darker than normal, e.g., normally whitish calypteres may appear yellowish, or yellowish calypteres a deeper yellow. All such factors will usually be compensated for in an adequate series reared from the same nest, but any one of them may seriously affect identification of single individuals.

Small size affects other characters. In small adults, the bristles are more slender and thus appear less well developed than in other species, even to the extent that ordinarily small bristles may appear to be absent altogether because they are hairlike and hardly distinguishable from surrounding hairs. For example, distinct accessory notopleural bristles are absent in the tiny specimens of braueri that were originally described as hirudo var. parva, although they are regularly characteristic of braueri. Undersized specimens may also yield atypical proportions. Because so few examples are available, two small females of eastern braueri are included in the measures for that species. Their ratios are the lowest in five of the six proportions used and obviously affected the ranges. For some proportions the ratios are extremely atypical.

Hall (1948) in his revision of the Nearctic species relied a great deal on measurements and ratios, but we find that his key is almost impossible to use because of the slight differences between species that are closely related and difficult to distinguish. For example, in his couplet 23 (his p. 101), he separated hirundo from sialia and metallica on the ratio of frons to head width, 0.09 vs. 0.10 or more in males, 0.27 vs. 0.28 in females. Moreover he stated that "Proportional measurements in specimens of the same species remain nearly constant." However, after hundreds of measurements, we find that there is a wide range in most ratios (cf. Tables V, VI) and that mean or median ratios can only be used effectively in separation of species where the differences are considerable. Standard deviations are of limited use where there are extensive overlaps of ranges. The range of ratios must obviously be considered in judging the usefulness of the mean ratio for a species, although extremes of ranges often represent one or few far-out specimens that would not seriously affect the identification of a great majority of specimens. The greatest differences in mean ratios often exist between species that are also distinct in some outstanding features that are easier to use, as for example the differences in male terminalia between avium and sialia.

Hall measured few specimens, five per species, or ten at most, and he included "the largest and smallest individuals,

Taxonomy of Adults 61

specimens from the extremes of the distribution, and several of average size," a mixture that would have greatly affected his ratios, especially in a statistical sample of few specimens. We differ in not measuring the smallest individuals, because the underfed "runts" in a series may and undoubtedly do show measurements and proportions that are not typical of the species. In our opinion, there is a normal or optimum size for each species, and measurements of such specimens will give a truer basis for comparison of species identities.

A random selection that included teneral specimens would give some distorted measurements. Selection must be confined to mature and measurable specimens, after which selection can be random to assure representation of the geographic range of the species. Even where it was necessary to measure a number of specimens from the same nest, presumably siblings, there was a range of variation and not the "nearly constant" measures postulated by Hall.

Accordingly the descriptions have been confined to a relatively few characters in which the mean ratios show some appreciable differences, and hence promise demonstrable significance as well as practicable usability in dealing with specimens. The best and most useful characters were found to be ratios between certain relatively stable features. These are listed below, with notes on how the measures were made, in order to promote uniform treatment and interpretation (cf. also Figs. 2a, b).

Width of frons: Measured across the narrowest part. In males this is usually anterior to the median ocellus, and in females at the vertex and measured across the posterior ocelli.

Width of head: The maximum over-all width, measured from the extreme convexity of each eye, as viewed directly from above. Special care must be given to the angle of vision, to assure the position for getting the maximum width.

Ocellar span: The width of the ocellar tubercle, measured between the outer margins of the posterior ocelli.

Width of parafacial: Measured opposite the anterior corner of the frontal vitta, at right angles to the eye margin and just below the foremost frontal bristle.

Vibrissal interval: The distance between the vibrissae, as measured between the inner margins of the two vibrissal sockets. This may not be too reliable. The epistoma often appears wrinkled or somewhat folded, or even concave, as if it is one of the last areas to become fully extended and hardened, and this could affect the measure and ratios based upon it. In some

62 Protocalliphora

measured specimens, the vibrissal interval was not measured because of this suspicion. The vibrissal interval may be characteristic of a species, but the practical usefulness of the character is limited by the degree of maturity of the specimen, often a serious problem with so many reared specimens.

Length of eye: The maximum length that can be observed, usually at an angle somewhat above direct profile. The maximum length is usually greater than its length as seen in direct profile, although the difference is not as great as in measures of the cheek height and does not seriously affect the proportion of cheek height to eye height (length).

Height (breadth) of cheek: The maximum height, measured at a right angle to the surface of the cheek, from a median point on the oral margin, midway between the vibrissa and the back of the oral cavity, to the lowest point on the eye margin. The height of the cheek compared with the length of the eye is a useful ratio, but the angle of measurement makes considerable difference in the cheek measurement and therefore in the ratios based upon it. This has been discussed by Sabrosky (1955) in connection with the tachinid genus Archytas. In many muscoid flies, such as Protocalliphora and Archytas, the cheek slopes or curves ventro-mesad from the eye, and its maximum height, measured at right angles to its surface, is somewhat greater than the height as seen in direct profile.

Features characteristic of one sex only are so specified, except that features found only in males (aedeagus, surstylus) are left unspecified. For other characters, if there is no qualifying note a statement applies to both sexes.

In the description of the male, the width of a parafacial is sometimes compared with the breadth of the 3rd antennal segment, but no figures and mean ratios are given. This is a relationship that can easily be judged by eye, but is not so easily measured, and it is used for gross comparisons, such as "obviously wider than" compared to "equal to."

Chaetotaxy: The chaetotaxy is rather uniform throughout the genus. The most notable variations are absence of reclinate orbital bristles (frontal bristles of Gregor and Povolny, 1959, and Peus, 1960) in females of braueri, presence of an accessory notopleural bristle (a smaller bristle between the two strong notopleural bristles) in that same species (Fig. 3a), and presence of two (occasionally three) posterior bristles on the fore tibia of chrysorrhoea and predominantly in tundrae and sapphira. Occasional individuals of other species, especially hesperia, show a second posterior bristle on the fore tibia on at least one side,

Taxonomy of Adults 63

and, less commonly, an accessory notopleural bristle may be seen. A distinct tympanic tuft of setae (Fig. 3b) is characteristic of chrysorrhoea and sapphira, but a few tympanic hairs occur occasionally in individuals of other species, and sometimes (e.g., braueri) in a considerable proportion of the individuals of a species. Variation in these characters is noted elsewhere, in the discussion of the species and in the discussion of the status of Apaulina Hall.

Male terminalia (Figs. 5-32): Examination of the male terminalia is usually essential for precise identification, and is always desirable. For some species, it is virtually the only significantly differentiating character, and species that are externally indistinguishable in the male sex may have quite different terminalia. Protocalliphora deceptor is the most startling example, the type series having been misidentified originally as a variety of aenea, until the terminalia revealed broad surstyli (Fig. 9) like those of avium. A few species can be identified without recourse to the terminalia, but even in those cases the terminalia can often confirm the identification. Frequently they will give an immediate clue to the identity, when variation is confusing or a teneral condition discourages measurement. The male terminalia are easily pulled into an extended position for study, even in dried specimens that have been relaxed over moist sand, although very old specimens will be difficult. For a discussion of technique and an explanatory figure see Hall (1948:37-39). In the present revision the "inner forceps" and "outer forceps" of Hall are termed cercus and surstylus. Past revisions of Protocalliphora, especially in Europe, have often showed the terminalia in posterior aspect. Unfortunately differences between the cerci of different species are usually slight, if any, and the posterior aspect misses the characteristic shape of the surstyli as seen in_ profile. Accordingly for most species we have figured a surstylus in side view to show how these differ greatly and how useful they can be in the differentiation of the species (Figs. 7-32). The aedeagus is fairly uniform in the subgenus Protocalliphora, differing from that in the subgenus 7rypocalliphora (Fig. 6a-c).

The female terminalia showed only slight differences, and these chiefly in species that were already amply distinct on external characters.

Size: In species reared from large species of birds, the young of which remain in the nest for a relatively long time, there is usually little difference in the size of the flies. Series reared from birds that develop quickly often show many undersized

64 —_ Protocalliphora

specimens, or "runts". Where material is adequate, the length is stated as a "normal length," 1.e., what appears to be the length normally attained by mature, fully fed individuals, but not including obviously immature or extremely undersized specimens. Most specimens of the genus are within 8-10 mm in length.

The sexes are described separately. The general matters of structure and chaetotaxy, described in detail for the male, apply also to the female unless otherwise noted. The normal differences between the sexes, e.g., the broader frons and the orbital bristles of the female, occur in all species and are not mentioned in the descriptions, except for the statements of ratios.

Most of the terms used are standard for flies (Diptera) and are explained and figured in the Manual of Nearctic Diptera, volume 1 (McAlpine et al. 1981), and usually in textbooks of entomology. The terms and measures used in this work are illustrated, somewhat diagrammatically, in Figures | - 4. Only a few terms need special mention:

(1) Accessory notopleural bristles (Fig. 3a): On each notopleuron in P. braueri, and only occasionally on other species, one to several bristles shorter and weaker than the standard 1 + 1 notopleural bristles or macrochaetae, but still clearly bristlelike compared with ordinary clothing hairs.

(2) Microtomentum (microtomentose): The fine coating or "dusting", called pruinosity in the Manual of Nearctic Diptera or pollinosity by some authors, is actually composed of ultramicroscopic outgrowths of the cuticle (Sabrosky 1983). These are cylindrical to more or less flattened and scalelike outgrowths, which grade from minute and fine to distinct and coarse. For descriptive purposes the term microtomentose can be appropriately modified, if desired, as finely, coarsely, thinly, or densely.

(3) Postalar wall (Fig. 3b): The vertical surface below the postalar callus. It may be bare or with few fine hairs, or with a tuft of coarse hairs.

(4) Preocellar area (Figs. 1, 2a): The part of the frontal vitta immediately anterior to the median ocellus. This may be a triangular to acuminate extension of the ocellar tubercle, especially in males, and dull or rugose or shining, or it may be an indefintely bounded polished spot, or it may be entirely dull and microtomentose, without any special spot or ridge or area, especially in females.

(5) Prevertical area (Fig. 2a): The narrow upper part of each parafrontal, just anterior to the vertical bristles.

Identification of Adults 65

(6) Surstylus (Figs. 7-32): In males, one of the clasping structures known by various authors as outer forceps, paralobes, or edita.

(7) Tympanic membrane or tympanum (Fig. 3b): An oval membrane, slightly countersunk in a "tympanic pit" behind the base of the wing and at the base of the calypteres, at the anterior end of the suprasquamal ridge. Like the postalar wall, it may be bare or with a few fine hairs, or with a tuft of long hairs (e.g., in chrysorrhoea).

Identification of the Adults

Few species can be identified satisfactorily in both sexes. In some species, males are distinct but females are not; in others, the revarse is true. In still other species, the immature stages are more distinctive than the adults. Crosskey (1981) has referred to a similar situation in the taxonomy of the Simuliidae: "Certain species in the better-known faunas can occasionally be quickly and easily identified on some distinctive feature ..., but it is unusual for a taxon to be equally distinct in the larvae, the pupae and both sexes of the adults."

The surest identifications of Protocalliphora depend, with few exceptions, on reared series in which both sexes are represented and puparia are associated. Accordingly the main key has been constructed on the basis that all three elements are present. The identity of the host may in some cases be helpful, or at least suggestive, but it is never by itself decisive. Lone specimens are extremely difficult to identify with any assurance, except for a very few unusually characteristic species, such as braueri and lata, or well-marked sexes such as males of deceptor.

The complex of species with curved surstylus could not have been solved without series of both sexes and their puparia, and identifications should be based on the same elements. When all these are available, some species can be spotted easily, such as brunneisquama with dark brown calypteres in both sexes, metallica and interrupta with greatly reduced spines on the puparium, metallica and bicolor with cupreous 5th tergite in females, and 5th cupreous also in male bicolor, and fallisi and tundrae with the posterior spine bands on the ventral side of the puparium vestigial or lacking. Recognition of these species narrows the problem to where, for the remaining species, ratios of measures can be utilized, variable though they are. However, when one tries to devise a key to males alone, the advantages of

66 — Protocalliphora

characters on the females and puparia fall by the wayside and, except for bicolor with its female-like cupreous 5th tergite, separation of species is much more difficult and the keys are less satisfactory. Likewise, for a key to females, once the cupreous 5th tergite has removed metallica, bicolor, and cuprina, identification of the remaining species is difficult or impossible.

We recognize, however, that it may sometime be desirable, for distribution or host records, to try to name isolated specimens, or a series composed of one sex only, or because the occasional mixture of species in a nest--12.1% in Bennett’s studies (1957), 7.1% in Whitworth’s (1976)--may be misleading and the sexes will require separate attention. (In all cases so far, one of the pairings, either of males or females or immature stages, alerted us to the existence of a mixture of species). Accordingly separate keys to males, females, and puparia have been prepared in which use of various characteristics, together with geographical distribution, will enable the user to eliminate some species and narrow the possibilities to one or a few names. In all three keys, after the especially characteristic species have been keyed out, discrimination between species becomes increasingly difficult because of variation in the few useful taxonomic characters, in the adults in the color of the calypteres, the number of posterior bristles on the fore tibia, and ratios derived from measurements. The key to males is more reliable than that to females because of the extra dimension provided by the male terminalia. Females of a number of species are "look alikes," and beyond couplet 12 that key is particularly weak. Puparia, except for a few relatively characteristic species, are difficult to key out by themselves, although in the main key they can be exceedingly useful in separating certain pairs of similar species, e.g., avium-asiovora, or metallica-cuprina.

In mixed infestations, the importance of careful association of each adult with the puparium from which it emerged is readily apparent. If puparia are available but not individually associated, they may do no more than reinforce a conclusion of a Oone-species series or of a mixed infestation, whichever is appropriate, although in some cases they may help to distinguish one or more of the species present, or to narrow the possibilities. In mixed infestations, more than any other, one must always be alert to the fact that not all collectors are careful to associate with an adult the very puparium from which that adult emerged.

In the separate keys to males and to females, puparia are sometimes cited for use when a series of reared specimens consists of one sex only, or on the chance that they might be

Identification of Adults 67

accompanying single specimens, although isolated specimens are most likely to be net-collected "loners."

Emphasis has been placed on characters that are easiest to use as well as most reliable. There was a temptation to begin with the choice of one or two posterior bristles on the fore tibia, the latter being one of the chief features of the species that may be termed a Palearctic or MHolarctic element in our fauna: chrysorrhoea, lata, and tundrae. However, a small number of specimens of species with a normal number of one posterior bristle have two on one or both fore tibiae, and some with a normal complement of two have only one, which would confuse anyone trying to identify specimens if that were the opening couplet (see notes on variation in tundrae and hesperia, for example). In a long series, one can determine the usual and presumably typical number, but this would not necessarily be true in every series. Consequently that character has been avoided as a primary choice, though mentioned as a supplementary character.

Species Groups in the Subgenus Protocalliphora

The species of the subgenus Protocalliphora can be arbitrarily divided into groups of convenience in various ways, depending on what characters are used, thus white calypteres vs. brownish, wide vs. intermediate vs. narrow frons in males (variable measures), monochromatic vs. dichromatic species, etc. The most useful division, and one that is especially good because the character states are distinct and can be illustrated, is based on the shape of the surstylus as seen in profile.

A. Surstylus broadly rounded (Figs. 7-9): asiovora, avium, deceptor B. Surstylus digitate (fingerlike or thumblike), parallel sided and straight, or nearly so (Figs. 10-19) Calypteres brown or brownish, especially margins: cuprina, halli, hesperia, hesperioides, lata Calypteres white: aenea, chrysorrhoea, hirundo, shannoni, sialia (yellowish in aenea)

68 Protocalliphora

C. Surstylus curved, usually slender (Figs. 21-32) Calypteres brown or brownish: brunneisquama Calypteres white:

Females with 5th tergite cupreous or cupreous green: bicolor, metallica

Female with bluish abdomen as in male: beameri, fallisi, interrupta, parorum, sapphira, seminuda, spatulata, spenceri, tundrae

Taxonomy and Taxonomic Characters of the Immature Stages

Chiefly because of the economic significance of the family, probably more attention has been given to the characters of the immature stages of the Calliphoridae than for any other group of the Cyclorrhapha, an effort directed primarily to easily available species. Only relatively recently, when more species were recognized in Protocalliphora, has greater attention been given to its immature stages. The 3rd-instar larva and puparium of Protocalliphora were first described and figured by Dufour in 1845 for what he called Lucilia dispar, now a synonym of P. azurea. By modern standards, this description was too generalized and incomplete. Hall (1948), while recognizing ten species of Nearctic Protocalliphora, gave descriptions of the immature stages of only two (3rd-instar larva of both, and puparium of one of those). A paper on the 3rd-instar larvae: of three European species was published by Rohdendorf (1957), well illustrated with figures of the important characteristics.

For the present study, early observations by D. M. Davies at Algonquin Park, Ontario, in 1949 were continued by the Bennett during 1950-56. Studies on the immature stages of the species in that area were broadened to include others, in order to parallel and complement the studies of the taxonomy of the adults being carried out by Sabrosky. A discussion of this joint study and an exhibit of both adults and immature stages were presented at the Tenth International Congress of Entomology in 1956 (Sabrosky and Bennett 1958). Studies since that time have added materially to the knowledge of the immature stages of the Nearctic species.

The present revision is fortunate in having material of the immature stages, in full or in part, of 23 of the 26 North American species recognized from adults (Table I). Puparia are known for 23 species and 3rd-instar larvae for 16 (plus two characters derived from the puparium for six others). As for the other stages, which are less useful, eggs are known for four

Taxonomy of Immatures 69

species, Ist-instar larvae for three, and 2nd-instar for seven. Long series, especially of puparia, were available for some of the species so that the extent of the variation could be assessed.

Preparation of Material

Treatment of material varied in detail from species to species and in decade to decade, but in general was as follows:

Second- and 3rd-instar larvae were split open from the prothoracic to anal segment along the lateral line (so as not to destroy the important diagnostic characters on the dorsal and ventral surfaces) and heated in a 5 molar potassium hydroxide (KOH) solution in small containers in a water bath. Heating continued until the sclerotized structures appeared to be adequately decolored, which varied from about 15 minutes to two hours. The larvae were then washed in hot water until all the internal structures washed free, leaving the cuticle with its attached spines, stigmatal plates and cephalopharyngeal skeleton. The larval skin was then cleared and dehydrated in beechwood creosote (which leaves the cuticle pliable) and mounted in balsam or "Permount", care being taken to ensure that the stigmatal area was folded to provide an unobstructed view of the stigmatal plates.

The puparia were heated for a variable length of time in 5 molar potash solution until adequately decolored and washed as above. The posterior region, portions of the dorsal and ventral cuticle, and the cephalopharyngeal skeleton (if present) were cut with a sharp scalpel from the pliable puparium, cleared and dehydrated in beechwood creosote and mounted as above. The procedure varied as conditions demanded. Certain puparia, such as those of P. aenea, with less color than others, required a shorter period of hydroxide treatment, while puparia such as those of P. avium and P. lata required much longer treatment. It was also true that old puparia, such as those from long-abandoned nests, required relatively little hydroxide.

All measurements are given as the average, the range (in parentheses) and the total number of measurements, e.g., 24 uw (14-41)/100. For spines, ideally ten measurements were made for spines on each portion of the cuticle for each individual puparium. When material was abundant, puparia were taken at random for measurements of length and breadth.

70 ~~ Protocalliphora

General Discussion of the Immature Stages

Several of the characters used in the following descriptions of Protocalliphora \arvae are not in common use in entomological literature, and therefore a discussion of the characters of the immature stages and a Glossary of terms are presented for the sake of brevity and clarity in the descriptions of species.

Glossary

anterior band: band of spines on anterior part of a segment’s ventral surface, posterad of a ventral pad.

anterior patch: dense patch of small, slender spines in the anterior band, immediately posterior to a ventral pad.

button (ecdysial scar): vestigial remains or scar from the shedding of a stigmatal plate of the previous instar, usually appearing circular or buttonlike at the base of the stigmatal plates.

cephalopharyngeal skeleton: the sclerites of the feeding apparatus of the larvae (mouth hooks, pharyngeal sclerites).

circumstigmatal folds: prominent ridges of cuticle surrounding and outlining the stigmatal area.

cuticular folds: prominent ridges of cuticle in the posterior region.

cuticular plaques: irregular masses of cuticle, heavily sclerotized, scattered throughout the cuticle. They are readily seen in the puparia and in stained preparations of the larvae.

cuticular ridges: ridges of cuticle, smaller than the cuticular folds, on the dorsal and ventral surfaces.

dorsal cuticle: cuticle of the dorsal surface of the larva and the puparium. Spines on the dorsal cuticle are usually directed posteriorly but in some species also anteriorly.

hyperstigmatal area: the area of cuticle immediately dorsal to the stigmatal area and bearing hyperstigmatal spines.

hypostigmatal area. the area of cuticle immediately ventral to the stigmatal area and bearing the hypostigmatal spines.

medial band: band of spines on medial part of a segment’s ventral surface, midway between the two ventral pads. The medial band is usually entire and pronounced, but in some species it can be markedly thinned or broken (interrupted).

mesostigmatal area: the area of cuticle between the stigmatal plates. It may bear mesostigmatal spines, folds and/or plaques.

parastigmate forms: puparia characterized by having the stigmatal plates abnormally close together as well as having

Taxonomy of Immatures 71

reduction in ratios and other features. Parastigmate forms are usually produced by under-fed larvae (see text).

peritreme: dark, sclerotized ring bounding each stigmatal plate.

plaques: see cuticular plaques.

posterior band: band of spines on posterior part of a segment’s ventral surface. It is usually a short arc of small spines immediately anterior to each ventral pad. It is sometimes absent in some species, and in others present only on the anterior five to six segments.

posterior region: at the posterior end (12th segment) of larva and puparium; that region bounded by a ring of posterior tubercles (cf. Hall, 1948, pl. 35D, E). The tubercles are fleshy and evident in the larvae, but are usually represented only by scars in puparia (especially in Protocalliphora).

prothoracic fringe (Hall 1948, pl. 38D): band of long spines on anterior margin of prothoracic segment of larvae and puparium. On ecloded puparia, it is often missing along with the ventral half of the puparial cap.

stigmatal area: broadly rectangular area surrounding and including the stigmatal plates; usually slightly depressed and frequently outlined by cuticular folds and/or small plaques. The area is usually divided medially by mesostigmatal folds and/or plaques, in some species bisected so sharply as to form two separate areas.

stigmatal plate (spiracular plate): each of two heavily sclerotized plates that bear two (2nd instar) or three (3rd instar and puparium) spiracular slits (stigmata); sometimes called spiracular plates.

stigmatal ratio: ratio of the distance between the buttons to the distance across the two stigmatal plates, measured between their outer margins at the level of the buttons.

ventral band ratio: ratio of the summation of the widths of the three ventral spine bands of any segment, measured on the midline, to the total distance between the adjacent margins of the ventral pads bordering that segment.

ventral cuticle: cuticle of the ventral surface of the larva and the puparium.

ventral pads (creeping welts): ventral, ovoid, fluid-filled, spine-free protruding areas of the larvae, situated on the midline in the intersegmental grooves, and apparently aiding in locomotion: comparable in position and function to spined, tuberculate, or roughened areas in other cyclorrhaphous larvae. The puparia show ovoid scars in the corresponding areas.

72 ~ Protocalliphora

Egg

The eggs are of typical calliphorid shape, rounded at one end and tapering slightly at the other, and lacking an operculum. The cuticle of the egg is variously ornamented with spines and raised ridges. Although eggs of only four species are described, the cuticle pattern offers specific differences, and examination of eggs of other species may also offer useful specific characters.

Larval stages:

First instar: The |st-instar larva differs, other than in size, from larvae of other instars as follows: (i) the lateral hooks and hypopharyngeal portion of the cephalopharyngeal skeleton appear to be elongated in comparison to those of other instars, but conversely the pharyngeal portion appears to be much shorter and the dorsal and ventral depressions are lacking; (ii) the prothoracic fringe is very short or absent; (iii) the spines of the anterior patch are two- to four-pronged in comparison to the simple spines of the other instars; (iv) mesostigmatal spines are absent; (v) hypostigmatal spines are scarce or absent; (vi) there is neither peritreme nor button.

Second instar. The spine distribution is the same as that of the 3rd instar, as is the general structure of the cephalopharyngeal skeleton.

Third instar. The tapering anterior end terminates in the cephalic segment which can be retracted into the prothoracic segment, on the anterior margin of which is a dense band of spines, the prothoracic fringe. These spines are longer than any Others on the body. The measurements of the spines of the fringe are based on the longest which are those of the posterior rows. In the formation of the puparium, the cephalic segment is withdrawn into the prothoracic, the most anterior external segment of the puparium. In this formation, the prothoracic fringe is contracted and the measurement of the fringe on the puparium is difficult, especially if the material is not well cleared in potash. Furthermore, in cyclorrhaphous flies, the prothoracic fringe remains on the lower half of the emergence cap of the puparium, and this section is frequently lost. The cuticular spines of the larvae are the same length as those found on the puparia. The cephalic annulus bears the oral opening through which the anterior portion of the cephalopharyngeal skeleton is

Taxonomy of Immatures 73

protruded. This skeleton, always present in the larva, adheres to the lower half of the emergence cap of the puparium and consequently is often lacking, as pointed out for the prothoracic fringe. Hence it is measured as from the larva, although the skeleton from the puparium is used in the absence of sufficient larval material. The skeleton was not found to be of use in the larval taxonomy of Protocalliphora, and only the overall length is given.

The key to the 3rd-instar larvae is based almost entirely on chaetotaxy, the lengths of the cuticular spines on various parts of the larva having considerable diagnostic importance. Characters such as the ventral band ratio, so useful in diagnosis of the rigid puparia, are not valid with the soft and pliable larvae, which, dependent on fixation technique, can be either contracted or extended.

Puparium

The color ranges from light brown in a thin-walled form to black brown in thick-walled species; puparia formed from underfed larvae are usually lighter in color. The surface is dull in the majority of the species, but appears shiny in the short-spined forms.

The larva will pupate either shortly after attaining the 3rd instar or after several more blood meals. In both cases, adults are produced, but the size range of both adult and puparium is considerable. Small larvae and puparia were found in nests shortly before the nestlings were fledged, and hence the size of the larva is most probably a function of the age of the nestling at time of attack. This is frequently the case with P. metallica, normally associated with ground-nesting sparrows and warblers, in which the nestlings are fledged in 8-12 days. On the contrary, in the nests of crows, hawks and other large birds in which the nestlings remain for 5-6 weeks, the puparia of P. avium are larger, on average, than any of the other species except P. lata. Length and breadth measurements are of limited use because of the degree of variation within a single species; breadth measurements given in text are taken at the widest point of the puparium.

The cuticle of both larvae and puparia of most species is thickly set with spines, the lengths of which vary in different regions of the body. The spines of any given region show a wide range in length, but the majority are of similar size. As more

74 Protocalliphora

puparia than larvae were available, the measurements of these spines were based on puparial material. The dorsal surface of the puparium is usually thickly set with spines, the largest in the center of the segment and the smallest towards the intersegmental annulus. The lengths of spines given in text are based on spines from the center of segment and these spines are termed dorsal cuticular spines. Shrinkage of the cuticle in the formation of the puparium results in the formation of numerous cuticular ridges, seen clearly on the dorsal and ventral cuticule. The size of these ridges seems to be constant for each species although the number found is variable; they are termed pronounced (sialia, Fig. 61d), weak (asiovora, Fig. 43b), or absent (aenea, Fig. 42d), according to size.

The ventral surface of the puparium presents a number of diagnostic features. A small vesicle, fluid filled in life, occurs on the midline of the intersegmental annuli of the larval stage. These vesicles, described by Guberlet and Hotson (1940), apparently serve the larva as a locomotory apparatus and here are termed the ventral pads (see Glossary). In the puparium, the remains of each ventral pad appear as a crescentric or ovoid, spine-free scar between segments (Fig. 43c,d). Between two adjacent pads each segment has, typically, three bands of spines. The degree of development in width of each band varies with the species. In the anterior band, immediately posterior to the ventral pad, is a dense patch of small, slender spines, termed the anterior patch. The second or medial band of spines, composed of stouter spines than those of the anterior patch, may be pronounced (Fig. 43c) or reduced (Fig. 50c). The third or posterior band of spines likewise may be pronounced (Fig. 44c), reduced (Fig. 43c), or absent (Fig. 42d). The degree of spination is expressed by the ventral band ratio, which is a decimal fraction derived from the sum of the widths of the three bands of spines on any segment (measured on the midline) divided by the total distance between the adjacent margins of the ventral pads bordering that segment. In general, and where possible, three or four ratios per puparium were calculated, based on measurements of the central and most typical segments 5 through 8. In samples with few available puparia, all were used but in large samples this was not the case.

The posterior region of the puparium has a number of useful characters. The sclerotization of the larval cuticle fixes the stigmatal (spiracular) plates, allowing certain measurements to be made. The diameter of a stigmatal plate is given as a maximal diameter measured from the dorsal portion of the peritreme

Taxonomy of Immatures 75

through the central spiracular slit to the ventral margin of the button. The distance between the two buttons (a) is also given, as is the width across the stigmatal plates at the level of the buttons (b), the measurements combined as a/b to give the stigmatal ratio. The peritremes of the puparia, unlike those of the larvae, are always closed. The cuticle between the stigmatal plates is termed the mesostigmatal area. Mesostigmatal folds and plaques are present or absent, depending on the species; if present, they bisect the area distinctly (bicolor, Fig. 45b), weakly, or not at all (Fig. 43a, b). The cuticle immediately dorsal to the stigmatal plates is termed the hyperstigmatal area, and with the exception of the prothoracic fringe, the spines of this region are usually the longest of the puparium. The cuticle immediately ventral to the stigmatal plates, the hypostigmatal area, characteristically possesses a number of folds; the number, size and configurations of these folds vary with the species. There are generally few spines in the hypostigmatal area compared to the number on the rest of the cuticle. The circumstigmatal folds pass laterally and dorsally from the hypostigmatal area around the stigmatal region; their formation, number and size vary with the species. Four basic patterns occur: irregular ridges of no fixed pattern (Fig. 44a); concentric ridges about the area (Fig. 57a); rectangular folds passing laterally but not surrounding the area (Fig. 56b); absence of folds (Fig. 43b). The basic patterns tend to be constant although the number of folds varies markedly; some species show more variability in these characters than others.

The posterior tubercles in the puparia are small scars surrounding the posterior region. They are not pronounced in most Protocalliphora (but more so on puparia formed from underfed larvae) and are more distinct on species with few cuticular folds. They are of little value taxonomically.

Production of small puparia by underfed larvae is frequent. These puparia tend to differ from normal puparia in the following respects: (a) the stigmatal plates are close together, forming a parastigmate puparium; (b) mesostigmatal characters are reduced or lost, consequent upon the juxtaposition of the plates; (c) circumstigmatal folds are frequently missing; (d) the ventral cuticle is compressed and the ventral spine bands are elided or otherwise distorted; (e) the cuticular spines are set closer together, giving the illusion of greater numbers and length; (f) cuticular ridges are usually absent; (g) posterior tubercles are usually pronounced.

76

5.

Protocalliphora

Key to the Nearctic species of Protocalliphora (based on males, females and puparia)

Male with unique surstylus and aedeagus (Figs. 6c, 20); female lacking upper (reclinate) orbital bristles; one or more accessory notopleural bristles present (Fig. 3a); puparium appearing smooth, cuticular spines small and scarce (Fig. 46), and prothoracic fringe extremely reduced, appearing absent .................00-

Peek t Maer, P. (Trypocalliphora) braueri (Hendel)

Male genitalia not as in braueri; female with reclinate orbital bristles (Fig. 1); accessory notopleural bristles rarely present (Protocalliphora, typical subgenus); puparium (except in two species) finely to coarsely spined, not appearing smooth, and with obvious prothoracic fringe.2

. Abdomen of males and females not concolorous, males blue

to bluish green or bluish purple (except in bicolor), females with 5th tergite cupreous, or abdomen almost

entirely aeneous to cupreous................. 3 Abdomen of males and females concolorous, metallic blue to DIUISH ee Cee se ny ec CNC tee Te 6

Female: thorax dark blue in ground color, abdomen blue or greenish blue with Sth tergite cupreous or at least cupreous tinted (occasionally 4th also); male: surstylus slender or digitate (Figs. 16,21,29) ............

Female: thorax and abdomen almost entirely aeneous to almost cupreous green; male: surstylus broadly digitate (Pigs TQ) ge Ons Ge as ee Se P. aeneaS. & D.

Abdomen of male blue, of female blue with Sth tergite cupreous (occasionally 4th also, see Sa, cuprina var.); male: surstylus digitate, or only moderately slender (Figs. 61, 21); puparium spined or not .............. 5

Both sexes with 5th tergite cupreous (sometimes weakly so if teneral) male: surstylus long and slender, slightly curved (Fig. 29); puparium rather strongly spined (Fig. 45) seastermise beans Steel Wave ee P. bicolor, n. sp.

Calypteres white in both sexes; male: surstylus of moderate width, slightly curved, parallel sided (Fig. 21); puparium with most spines present only as minute tubercles, appearing like fine sandpaper (Fig. 56); chiefly eastern

dea se cates few Var Sen Ne nea asa areola P. metallica (Tns.)

Key to Nearctic Species 77

Calypteres brownish in male, white in female; male: surstylus digitate or only slightly curved, broader than in metallica (Fig. 16); puparium with distinct though short spines (Fig. AS) si western oN Ce, Bae I, P. cuprina (Hall)

. Parafacial unusually broad, over three times breadth of 3rd antennal segment; calypteres dark brown; male: frons unusually broad, at narrowest equal to or even greater than width of an eye, and with both proclinate and reclinate orbital bristles (Fig. 33); puparium strongly ridged and spined (Fig. 55); western ... P. latan. sp.

Parafacial narrower, at most barely twice breadth of 3rd antennal segment; calypteres unusually whitish, brown in only a few western species; male: frons at narrowest obviously much less than width of an eye, usually much less, always lacking orbital bristles (Figs. 34-38);

puparium variable, usually not strongly ridged .... 7 . Male: surstylus broadly rounded (Figs. 7-9) ........ 8 Male: surstylus digitate to slender (Figs. 10-32) ...... 10

. Male: frons strongly narrowed, narrowest width averaging 0.07 times width of head and obviously narrower than breadth of 3rd antennal segment, parafrontals almost touching; female: frons narrow, half or less the width of an eye, averaging 0.23 times width of head; puparium with minute spines, the appearance like fine sandpaper or little more (Fig. 49); eastern ...... P. deceptor n. sp.

Male: frons broad for the genus, range 0.115-0.18 times width of head and obviously much greater (about twice) than breadth of 3rd antennal segment, the parafrontals widely separated (Figs. 34, 35); female: frons broad, obviously over half width of an eye and averaging 0.295 times width of head; puparium well ridged and spined (Pigsv4 se 44) as a Me ee MN, 9

. Male: frons typically wider than in asiovora, averaging 0.16 times width of head (Fig. 34); puparium with medial spine band of ventral cuticle pronounced and uninterrupted (Fig. 44); eastern ... P. aviumS. & D.

Male: frons narrower than in avium, averaging 0.13 times width of head (Fig. 35); puparium with medial spine band of ventral cuticle pronounced but distinctly interrupted on midline (Fig. 43); western .... P. asiovoraS. & D.

Protocalliphora

. Postalar wall and tympanic membrane each with conspicuous tuft of numerous long, brown to blackish hairs (Fig. 3b); fore tibia typically with two posterior bristles; almost always in nests of bank swallow ...............

IO RE ile ca a a meer a Ree P. chrysorrhoea (Meigen) Postalar wall and tympanic pit bare or with one to few, usually short and inconspicuous, usually pale hairs; fore tibia with one posterior bristle in most species, commonly two in sapphira and tundrae; rarely in nests of bank SWAllO we icici, ie ON HO athe Gr ical ca cans ha 11

Calypteres brownish or brownish yellow in males, same in females (except in halli), sometimes paler brown but at least brownish tinted with outer rims and fringe brownish to brownish yellow; western species. ........... 12

Calypteres in both sexes white ......555......054- 15

. Surstylus in male exceptionally long and slender, curved (Fig. 28); parafacial obviously much wider (nearly twice) than breadth of 3rd antennal segment ................

SE eA e SNM Cosa Dr OL WBN ae Malan P. brunneisquama, Nn. sp. Surstylus shorter and/or broader, not or only slightly CURVER: PEEVE Re INGE ST En Shai hoe 13

. Calypteres brownish or brownish yellow in both sexes . 14 Calypteres brownish or brownish yellow in male, white in PEMA ee) Bee Sern ise re cak eae hee alee ee an P. halli, n. sp.

Preocellar area in female heavily and evenly brownish microtomentose, in male almost always gray and narrowly triangular, rarely somewhat shining; palpi black-tipped

SANE HRC TOT Ty ene ry coir brake Oe ted P. hesperia S. & D. Preocellar area in female polished black immediately anterior to median ocellus, in male with narrow polished line or

low ridge (as in Fig. 2a); palpi usually entirely yellow RNA LSE amie oma A Oh CORN cp P. hesperioides, n. sp.

. Surstylus digitate, not appreciably curved, and broader than opposing species (Figs. 13-15) .............. 16 Surstylus distinctly curved, usually slender on distal half or more, or parallel sided throughout (Figs. 22-32) . 19

. Frons of male obviously wider than breadth of 3rd antennal SCE MENGE 35s onside a SAR ead ee core 17

Key to Nearctic Species 79

-- Frons of male obviously narrower than breadth of 3rd antennal segment ..;.......... P. shannoni, n. sp.

17. Male with polished black and narrow triangle or streak anterior to median ocellus (Fig. 37); preocellar area in female shining or gray; prothoracic fringe of larva or puparium long and dense, spines averaging 150 in lensthe(SO= 200) ii 395 ce Se erm eee) ae 18

-- Male with preocellar triangle gray microtomentose (Fig. 38); preocellar area in female entirely gray microtomentose; prothoracic fringe not as conspicuous, spines averaging 57

in length (21-103) .......... P. hirundo S. & D.

18. Preocellar area polished black in both sexes; eastern ... ich a ME Oe: UAE QSL Ca Seth a TT I P. sialia S. & D. -- Preocellar triangle or streak polished black in male, but preocellar area dull and heavily microtomentose in female; western ........... P. sp. (western sialia)

*19. Puparium: hyperstigmatal area and dorsal cuticle with GistinctySspimes 3 eee VG. Lee ea ke 20 -- Puparium: Spines of both hyperstigmatal area and dorsal

cuticle minute (fine sandpaper appearance) (Fig. 54) . Be HEDIS R NE SEE MR Ve NAN TVD Bile bea ee att P. interrupta, n. sp. *N.B. The puparium is presently unknown in beameri and sapphira but assumed to have distinct spines; if not, in both of these the frons is relatively broad compared with that in interrupta (0.12-0.17 times head width compared to 0.09 in interrupta). If teneral or incompletely colored metallica slipped through couplet 3 and came to this point, its surstylus

is broader than in interrupta (cf. Figs. 21 and 30).

20. Parafacial in both sexes relatively broad, obviously wider than breadth of 3rd antennal segment ......... 21

-- Parafacial in both sexes narrow, equal to or barely wider than breadth of 3rd antennal segment P. spenceri, n. sp.

21. Male with preocellar area broadly triangular, evenly bright gray microtomentose (Fig. 36); fore tibia typically with two posterior bristles; far north, arctic Canada and Greenland. oie en P. tundrae, n. sp.

-- Male with preocellar area narrowly triangular to acuminate, often more or less rugose and shining; fore tibia with one posterior bristle in most species; not far northern .. 22

80 Protocalliphora

22. Male with occiput predominantly white haired behind and below the row of ostocular setae, at most only a few slender black hairs in a partial row P. beameri, n. sp.

-- Male with numerous coarse black hairs in two or more rows behind and below the postocular setae ........ 23

23. Frons of male heavily setose, frontal bristles and parafrontal hairs extending posteriorly at least to level of ocellar bristles; parafrontal relatively wide all the way to VEOTCOX HE I Oh is NUN, eee ai mY Raleok P. spatulata, n. sp.

-- Frons of male seldom as setose as in spatulata, frontal bristles and parafrontal hairs usually ending anterior to level of median ocellus; parafrontal strongly narrowed dorsally in Preventical area ceca EO SOR es ee aire 24

24. Frons of male narrow, averaging 0.08 times head width (combined range of three species 0.06-0.13); parafacial of male wider than frons at its narrowest; fore tibia with one posterior bristle... 20k ech Oes Cie as 25

-- Frons of male broader, 0.17 times head width in the one known male; parafacial of male narrower than frons (OLS 5S) ce See SUA ae eek nh. ain P. sapphira (Hall)

25. Parafrontals of male wide, each wider midway than narrowed frontal vitta: parafrontal and parafacial of male sparsely and finely haired, contributing to an "almost bare" appearance, silvery microtomentose ............

pH SEEN Sed Rule SIU EL eReEe eu Si P. seminuda, n. sp.

-- Not as above, frontal vitta of male much wider than a

parafrontaleid) cheer eee eile, Gee Ene tee 26

26. Puparium ventrally with posterior band of spines well developed, thus three spine bands per segment (Fig. 57d); head dark gray microtomentose; typically parasites of chickadees (Parus spp.); western .. P. parorum, Nn. sp.

-- Puparium ventrally with posterior spine band vestigial or lacking on at least posterior five or six segments (Fig. 50c); parafrontal and parafacial brighter, silvery gray microtomentose; typically parasites of marsh-inhabiting birds; northeastern’. 22). P. fallisi, n. sp.

6.

Key to Males 81

Key to males of Nearctic Protocalliphora

. Aedeagus unique in genus (Fig. 6c); one or more accessory

notopleural bristles (Fig. 3a); frons extremely narrow (Fig. 41), averaging only 0.05 times width of head and less than half breadth of 3rd antennal segment, at narrowest parafrontals almost touching ................... SEA Lee Reap P. (Trypocalliphora) braueri (Hendel) Aedeagus not as in braueri (Fig. 6a,b); usually no accessory notopleural bristles; frons usually broader and parafrontals well separated (Figs. 33-40) (typical subgenus PTOLOCQUIIUPNOTQ) ae ook RM aN ot ted DE

. Calypteres deep yellowish to brown or brownish yellow 3

Calypteres white or whitish, at most outer margins pale VOLE We) ROLL G eS RCO UR CAAT ARON IONE cou aS Oe 9

*N.B. P. aenea, usually deep yellowish, is keyed both ways because of possible misinterpretation. It is somewhat intermediate, especially if individuals are immature and not fully colored.

(Calypteres deep yellowish to brown)

Frons unusually broad for a male, at narrowest equal to or even broader than width of an eye (Fig. 33); proclinate and recinate orbital bristles present, as in females; parafacial unusually broad, over three times breadth of 3rd antennal segment; calypteres dark brown......

Aryan bait hig) ee taliy SARL GY 0 ee ibe aha P. lata, n. sp.

Frons obviously much narrower than width of an eye; no

orbital bristles; parafacial obviously not as broad... 4

. Calypteres yellow to deep yellowish ... P. aeneaS. & D.

Calypteres definitely brown to brownish, at least rims and EMPRIN GC Sieg ie eo ce Vee Tee a MAAN CAI UNNI Ga 5

Surstylus exceptionally long, slender, and curved (Fig. 28) AE IN ye als oh Ua eet CC SO EN P. brunneisquama, Nn. sp. Surstylus digitate, not curved, straight or nearlyso ... 6

Palpus entirely orange-yellow; preocellar area a narrow polished line or low ridge (Fig. 2).............. SS ie gn PERN ca al Ua Ana dD ee P. hesperioides, n. sp.

1

Protocalliphora

Palpus black-tipped; preocellar area usually narrowly triansularto,acumimate: 2) 3 4 isda es oa ee 7

. Preocellar area usually triangular and_ evenly

gray-microtomentose ............ P. halli, n. sp. Preocellar area narrow and acuminate, more or less rugose and shining or subshining................... 8

Parafacial relatively narrow, equal to or barely wider than 3rd antennal segment ........... P. cuprina (Hall)

Parafacial obviously wider than 3rd antennal segment 2 Ae Wabssiro anus iat ule) (aie aS ear aa P. hesperia S. & D.

(Calypteres white)

. Surstylus unusually broad (Figs. 7-9) ............ 10

Surstylus slender to digitate (Figs. 11, 13-15, 21-32) . 12

. Frons strongly narrowed, averaging 0.07 times width of

head, obviously narrower than 3rd antennal segment, and at narrowest parafrontals almost touching; cerci in posterior view narrowly acuminate, parallel to acute apices (Big. Sb) igs a ee P. deceptor, n. sp. Frons broad, averaging 0.13-0.16 times width of head and obviously much wider than breadth of 3rd antennal segment, parafrontals widely separated; cerci sturdy, divergent, and apices thick and rounded (Fig. 5a) 11

Frons typically wider than in asiovora, averaging 0.16 times width of head (Fig. 34); surstylus narrower in proportion to its length than in asiovora (Fig. 7); eastern .....

EDT AC ROR NSE AL SMe TU a Nha CaM Mtn chai T P. avium S. & D.

Frons narrower than in avium, averaging 0.13 times width of head (Fig. 35); surstylus broader in proportion to its length than in avium (Fig. 8); western ............

Sie sabi Recetas eae taza nteiee SN P. asiovora S. & D.

. Fifth tergite cupreous (sometimes weakly so, if teneral)

Pepe neE er iea SA TE MIB LER an) Ceara LEN P. bicolor, n. sp. Fifth tergite blue to bluish purple ............... 13

. Surstylus digitate, not appreciably curved, and broader than

opposing species (Figs. 11-15) .............. 14 Surstylus distinctly curved, usually slender on distal half, or parallel sided throughout (Figs. 21-32) ........ 18

14.

18.

Key to Males 83

Postalar wall and tympanic membrane each with conspicuous tuft of long, brown to blackish hairs (Fig. 3b); fore tibia typically with two posterior bristles ............

Hy 2 Any RR a ADRS Oe LEE Ean ae P. chrysorrhoea (Meigen)

Postalar wall and tympanic membrane with one to few, usually short and inconspicuous, usually pale hairs; fore tibia with one posterior bristle in most species... 15

. Frons obviously narrower than breadth of 3rd antennal

SESE NCE) i ee Sia th GaN eR TG epee ee So eral Fae 16 Frons obviously wider than breadth of 3rd antennal segment RU MERE CSAS GN inh rte IM: eae Rehab ehh. battens, 17

. Calypteres always at least slightly yellowish, with dark yellow

margins (Typical males of aenea have brownish yellow calypteres with brown margins) ... P. aeneaS. & D.

Calypteres white with pale yellow margins .......... AT RE oars Pee ocean a Pie RAG: P. shannoni, n. sp.

Preocellar area with polished black and narrow triangle or low ridge (Fig. 37); surstylus in profile relatively long and digitate (Figs IS\ie 8 sae Ae: P. sialia S. & D.

Preocellar area with broad and evenly gray-microtomentose triangle (Fig. 38); surstylus in profile appearing stubby, relatively short for its length and broad at base (Fig. MA area tir iat Co cl a te etna aie ce eect P. hirundo S. & D.

Preocellar area broadly triangular, evenly bright gray microtomentose (Fig. 36); fore tibia typically with two posterior bristles; far morth, arctic Canada and Greenland: osc. il ee P. tundrae, n. sp.

Preocellar area narrowly triangular to acuminate, often more or less rugose and shining; fore tibia with one posterior bristle in most species; not far northern ....... 19

Frons heavily setose, frontal bristles and parafrontal hairs extending posteriorly at least to level of ocellar bristles; parafrontal relatively wide all the way to vertex . 20

Frons seldom as setose, frontal bristles and parafrontal hairs usually ending anterior to level of median ocellus; parafrontal strongly narrowed dorsally in prevertical ATCA Meee ALC aN) eee Cosa cles CREE Man anes LAER chee 21

84

20.

PANN

Protocalliphora

Surstylus strongly narrowed on distal half and shorter than céren(Figh 31) sci oe is ae P. beameri, n. sp.

Surstylus parallel sided for most of its length, usually

widening rather abruptly at distal end to appear spatulate (Bigs 27) hy eae ee P. spatulata, n. sp.

Parafacial wider than width of frons at narrowest; frons averaging 0.08-0.12 times head width; fore tibia usually with one posterior bristle .................. 22

Parafacial narrower than width of frons (0.85x); frons 0.17 times head width in the one available specimen; fore tibia with two posterior bristles (probably variable) .....

Bag 5 sinensis) /al Cain TREN rear ea her AAG Ot RT P. sapphira (Hall)

22. Parafrontal and parafacial dark gray microtomentose;

24.

—

typically parasites of chickadees (Parus spp.); western GE ERR BROS ae ar ah eR P. parorum, n. sp.

Parafrontal and parafacial bright, silvery gray

MICFOtOMENTOSE EH 8 ee BO eee

Parafrontals relatively wide, each wider midway than narrowed frontal vitta; parafrontal and parafacial sparsely and finely haired, contributing to an “almost bare" appearance, silvery microtomentose P. seminuda, n. sp.

Not as above, frontal vitta much wider than a parafrontal

BU ESTEE MU Lao, ter ellis cane Saka ee a ee 24 Parafacial opposite lunule broader than 3rd antennal SESMEMES PSU eS CNN An EN eae ounce 25 Parafacial narrower than breadth of 3rd antennal segment, or barely equal to it ........... P spenceri, n. sp. SOWOSEORIN Se eta entice ound nen onl ane P. interrupta, n. sp. Eastern co ch ere ee Ce aaaeay aa P. fallisi, n. sp. Chiefly eastern sy es eas P. metallica (Tns.)

Key to females of Nearctic Protocalliphora

. Upper (reclinate) orbital bristle present; usually no accessory

notopleural bristles (P., subgenus Protocalliphora) . 2 *No upper orbital bristle; accessory notopleural bristles Present Ve eas P., subgenus Trypocalliphora)

(one species, P. braueri (Hendel))

Key to Females 85 *N.B. See Variation under P. spenceri, in which two females are aberrant in lacking upper orbitals, but accessory notopleurals are also lacking.

Thorax and abdomen almost entirely aeneous to almost

CUPREOUSTSFeeN GAS, Be ale sas P. aenea S. & D. Thorax and abdomen bluish to bluish purple or greenish blue, 5th tergite cupreous in three species ........... 3

. Calypteres brown or brownish (tending to be brownish yellow

inimmature Specimens) 2.3.0... 3k. Slee 4 Calypteres white or whitish, at most outer margins pale Ve llowaASh eis een Tice AUN ne Sm Reon away See ene co, il

*N.B. P. hesperia is keyed both ways because in some series females have pale calypteres, yellowish brown or even somewhat whitish, possibly because of immaturity and lack of full coloration in the reared series.

(Calypteres brown)

Parafacial unusually broad, over three times breadth of 3rd antennal segment; fore tibia with two posterior bristles; WESTON Riel ts RE oa tay SUSAR a3 P. lata, n. sp.

Parafacial narrower, at most about twice breadth of 3rd antennal segment; fore tibia with one posterior bristle in MNOSTESPSCISS oe sis 28F A Eh aR en Ria ego. 8) aye Seats 5

. Parafacial obviously much wider than (nearly twice) breadth

of 3rd antennal segment... P. brunneisquama, Nn. sp. Parafacial obviously much less than twice breadth of 3rd antennalisegment:is saci i aa a ene 6

. Preocellar area polished black, small but distinct (cf. Fig. 2a);

palpus entirely orange-yellow . P. hesperioides, n. sp. Preocellar area dull, heavily gray to brownish micro- tomentose; palpus black-tipped.. P. hesperiaS. & D.

(Calypteres white)

. Fifth tergite cupreous to cupreous green .......... 8

Fifth tergite bluish or greenish blue, concolorous with precedingstergites i000. ea a ear 10

86

12%

Protocalliphora

. Anterolateral area of scutum almost solidly shining

blue-black, as viewed from behind; if puparium present, most spines greatly reduced, minute, the appearance like that of fine sandpaper (Fig. 56) . P. metallica (Tns.) Anterolateral area of scutum usually not so, as viewed from behind, the shining area deeply incised by extension of posthumeral gray microtomentum, extreme in cuprina, weak to somewhat intermediate in bicolor; if puparium present, spines are obvious ................--

Western acer Sa ee eee es aaa 9a Easterns 28 etre er Aes ene ae ee. P. bicolor, n. sp.

. Only 5th tergite cupreous or cupreous green ........

Le Nae ia) a ee RU Lg 5. P. cuprina (Hall) Fourth and Sth tergites both cupreous or cupreous green Se aE hs an ey Nave P. cuprina var. or n. sp.

Frons exceptionally narrow, averaging 0.23 times width of head; parafrontal relatively wide and frontal vitta less than twice width of a parafrontal; eastern .........

PSI NN AU SSH IN oer SRN P. deceptor, n. sp.

Frons not so narrow and frontal vitta broader, two to three

times width of a parafrontal ................ 11

. Postalar wall and tympanic membrane each with conspicuous

tuft of numerous long black hairs (Fig. 3b); fore tibia typically with two posterior bristles ............

SUA TALENTS. SOME ei abaalNraua T0340 P. chrysorrhoea (Meigen)

Postalar wall and tympanic membrane bare or with one to

few, usually short and inconspicuous, usually pale hairs;

fore tibia with one posterior bristle in most species,

commonly two in sapphira and tundrae ........ 12

Scutum heavily microtomentose anterior to transverse suture; median shining stripe absent or indistinct as viewed from behind at a low angle, two narrow dark stripes outside acrostical bristles more distinct than usual, the resulting appearance being that of three broad gray stripes; WESTORT ois ue Eau ne P. spatulata, n. sp.

Scutum not so heavily microtomentose, usually with a broad median shining stripe flanked by broad gray microtomentose stripes, as viewed from behind at a low 11h: d (= ERDAS WOR Aa WAR SLR URe) Cael apne MUL LS Ri DE 13

20.

Key to Females 87

Eastern North America, arctic and subarctic Canada, and Greenland riecieenrnewad. oar iis ee Maa eee. a. 14 WesterhnsNorthpAmerica 127). (O8 ei eee ee 18

. Parafacial relatively broad, width at lunule obviously much

greater than breadth of 3rd antennal segment ... 15 Parafacial relatively narrow, barely wider than breadth of 3rd antennal segment .......... P. shannoni, n. sp.

. Arctic Canada and Greenland; typically with two posterior

bristles on fore tibia .......... P. tundrae, n. sp. Not far northern; typically with one posterior bristle on fore tibia: (twoun part of fallisi!)) se se. ws 16

. Parafacial about twice breadth of 3rd antennal segment,

usually slightly convex and appearing bulging; robust

species, normal length 9.5-10 mm.. P. avium S. & D. Parafacial not as broad, and not appearing bulging; smaller SDECIES = 77.-9=9 DM i A ee ia 17

. Preocellar area a large, irregular, polished black spot

CHG eH RR sme ac a es ne ge P. sialia S. & D. Preocellar area dull, finely and evenly microtomentose, sometimes thinly so and subshining ._ PP. fallisi, n. sp.

Dark species, head dark gray microtomentose; parafrontal posteriorly tending to be thinly microtomentose, hence

subshining, blackish........... P. parorum, n. sp. Brighter species, bright gray microtomentose, often tinted yellowishsor brownishic:, ies. S28 ke hc ees 19

. Preocellar area partly but irregularly polished black,

sometimes thinly microtomentose and _ subshining; parafacial unusually narrow, equal to or barely wider than breadth of 3rd antennal segment... P. spenceri, n. sp. Preocellar area evenly and usually heavily microtomentose; parafacial broader, usually obviously wider than breadth of 3rd antennal segment ..................-. 20

Fore tibia with two posterior bristles, perhaps typically, but probably variable; parafrontal, parafacial, and preocellar area decidedly brownish yellow .. P. sapphira (Hall)

88

2D

23%

Protocalliphora

Not with that combination of characters, fore tibia typically with one posterior bristle (variable in some species!), and parafrontal, parafacial, and usually preocellar area bright gray or light yellowish-gray microtomentose .... 21

. Parafacial relatively narrow, appearing equal to or only

slightly wider than breadth of 3rd antennal segment 22

Parafacial relatively broad, obviously much wider than breadth of 3rd antennal segment, usually almost twice

3 SUPRA SOROS, COT STIS es ROGAN Ge Nae ee 23

Females of three species cannot be reliably separated Re meaner ec mW pie cg P. halli, n. sp. PTET EU RET PGC Leg okt Mere UCN ages) ANN P. hesperia S. & D. Ree Uae Shea: AY SEES Henne P. interrupta, n. sp.

Females of five species cannot be reliably separated Rt REET ECE AN Rh PIR SRC a 8 P. asiovora S. & D. AAR Se aA CaS EU ae P. beameri, n. sp. Beal mR Bren tte Une back pale P. hirundo S. & D. ATI REL PL CE ISA ON PRD BEEN AK P. seminuda, n. sp. 2 ANG ESS IRENE es SU cin Morals MURA Hb Ah P. western sialia’

Key to the 3rd-instar larvae of Nearctic Protocalliphora

Prothoracic fringe much reduced (average less than 10. ); cuticular spines much reduced in number or absent, at most limited to a single narrow band per segment ...

A Ry P. (Trypocalliphora) braueri (Hendel)

Prothoracic fringe pronounced (averaging 25 or more); cuticular spines numerous on all parts of body (Protocalliphora, typical subgenus) ............

. Medial band interrupted on midline .............. 3

Medial: band’ entire jee Se) Te eee 4

. Dorsal cuticular spines small, averaging less than 10

SR Ne RA Re SN Ae P. interrupta, n. sp. Dorsal cuticular spines larger, averaging more than 15. RC eRe HE tas 1A minal mea les a aa P. asiovora 8. & D.

. Prothoracic fringe averages more than 100 _...............

ESF ucla, ual Wel aaa eee ree A Na A P. sialia S. & D.

Key to 3rd-Instar Larvae 89

. Hyperstigmatal spines as tubercles, averaging 4-6 wu .. 6

Hyperstigmatal spines as spines, larger than above ... 7

. Entire dorsal cuticle with spines reduced to minute tubercles;

spines of medial band average 6 (2-12) W ......... Rava tan mene ts ai ae mY ete P. metallica (Tns.) Dorsal cuticle with definite, although relatively few, dorsal spines; spines of medial band average 12 (2-29) uw... Lo SACRE OE eat UN UGe nan Maia MSD Dh pe ed ia P. deceptor, n. sp.

Posterior band vestigial or absent on at least posterior 5-6 SEQIMEMES aye a ON RUD UE RIA RS A NAVD AUR och 8 Posterior band present on all segments ........... 1]

. Posterior band reduced but present on anterior 5-6 segments

i ON CERCA MM oo OS) ire Te SERINE DR P. tundrae, n. sp.

. Prothoracic fringe averages more than 70 wu; hyperstigmatal

spines )34)((16=52)) pb Wee eas P. aenea S. & D. Prothoracic fringe averages less than 60 [; hyperstigmatal spines average less than 30 w ............... 10

. Hyperstigmatal and dorsal cuticular spines average 29 pw or

JUOVG) 2} bai eae calle ae me RET ea ba P. halli, n. sp. Hyperstigmatal and dorsal cuticular spines average less than oyR 74 AOD eM re yan me Pu din SATO eA WR TAN lh U4 P. fallisi, n. sp.

. Prothoracic fringe averages 80 / or more ........ 12 Prothoracic fringe averages 75 Wor less ......... 13

. Hypostigmatal spines average 6 uw .... P. bicolor, n. sp.

Hypostigmatal spines average 15 for more .......... ERAT ian aA NS eae ea eh P. parorum, n. sp.

. Prothoracic fringe averages 60 &@ or more ........ 14 Prothoracic fringe averages less than 60 w........ 17 . Prothoracic fringe averages more than 70 “w....... 15 Prothoracic fringe averages less than 70 @........ 16

. Spines of dorsal cuticle and medial band average more than

AOE a ae UNA at en) NN Va aI ea a ls P. lata, n. sp. Spines of dorsal cuticle and medial band average less than 30 TDC AD AL, die te ge eta aes Ue Magen ASI P. hesperioides, n. sp.

90 = Protocalliphora

16. Hyperstigmatal spines average 28 wu; hypostigmatal spines

average USL acai Cees ae P. cuprina (Hall) -- Hyperstigmatal spines average 11 wu; hypostigmatal spines as tubercles, averaging 6mw........ P. shannoni, n. sp.

17. Prothoracic fringe averages 29 (12-62) Ww ........... oe Ry Ne lata Meese De ane P. chrysorrhoea (Meigen) -- Prothoracic fringe averages more than 40 w....... 18

18. Spines of anterior patch small, average 9-12 Ww ....... SPIT HE IN BONE UT Ry cen a a eas P. *spenceri, n. sp. Bs RENEE SOE Ne MNS Se Maa P. *seminuda, n. sp. *N.B. Third-instar larvae of these species cannot be separated on the basis of larval characters alone. -- Spines of anterior patch averaging 15 wor larger.... 19

19. Spines of dorsal cuticle and medial band average 20 wp. SIA A RI Ne LENS AON SCURRY ibe P. avium S. & D.

)O0KO} Ca) Sts aid) oy oon ONG Ol OsOlOl Msu0.6 Olb1G ola Goo c0 04d o.6

*20. Posterior band pronounced; ventral band ratio more than OPS Or ey iain ee eee Nine aad Grenae P. spatulata, n. sp. -- Posterior band weak; ventral band less than 0.70 ..... Dy WARES eae ON a nia tt QiOrS MBL S MAN P. hirundo, S. & D. *N.B. Ventral band ratios in larvae are not reliable. Species in couplet 20 can only be identified with certainty on the basis of puparial or adult characters.

Key to the puparia of Nearctic Protocalliphora

1. Dorsal cuticular spines much reduced in number, at most limited to a narrow band per segment; prothoracic fringe vestigial, spines averaging less than 10 w..........

Se eats P. (Trypocalliphora) braueri (Hendel)

-- Dorsal cuticular spines numerous (in two species small or minutely tuberculate); prothoracic fringe well developed,

spines averaging more than 25 yp (Protocalliphora, typical Subgenus) :..)2 BUHL S eheD PEERS ei ee eae 2

2. Medial band interrupted on the midline ........... 3 ==) (Medial band entire... 0.) ek Ca eee 4

Key to Puparia 91

3. Dorsal cuticular spines large, averaging 25 (12-45) uw; ventral band ratio 0.63; posterior band pronounced .......

SE ORTEGA DEE OR UT ero Ome P. asiovora S. & D.

-- Dorsal cuticular spines small, many as tubercles, averaging

4-6 uw; ventral band ratio 0.27; posterior band usually

absent or at best, vestigial ..... P. interrupta, n. sp. 4. Hyperstigmatal spines as minute tubercles, 4-6 uw .... 5 -- Hyperstigmatal spines as spines, more than 1OW..... 6

5. Dorsal cuticle with spines reduced to minute tubercles, with a fine sandpaper-like appearance; spines of medial band average 6 (2-12) W........... P. metallica (Tns.)

-- Dorsal cuticle with definite spines, not appearing as fine sandpaper; spines of medial band 12 (4-20) uw .....

Pee iias einai Megs ubeella patiay Rela aM pe P. deceptor, n. sp.

6. Posterior bands vestigial or absent on at least posterior 5-6 segments; ventral band ratio normally less than 0.50 7 -- Posterior band present (usually pronounced) on all segments; ventral band ratio normally than 0.50 ......... 10

—~

. Posterior band present, although much reduced, on anterior 5-6 segments; circumstigmatal folds present ..... 8

-- Posterior band absent, or at best vestigial, on all segments;

circumstigmatal folds absent.................

oo

Posterior band, although reduced, present on anterior 5-6 SCOMENTS ees Ae IN eae ae, P. tundrae, n. sp.

-- Posterior band absent or vestigial on all segments aa

Seo natrrse Alien ticles Rae etna Rie ie aa a P. halli, n. sp.

9. Prothoracic fringe averages more than 70 yu ventral band ratio more than 0.40; dorsal cuticular spines large ......

TERMS Stet He Uy SOR A TCA arin tA ays SHARO P. aenea, S. & D.

-- Prothoracic fringe averages less than 60 wu; ventral ratio less

than 0.40; dorsal cuticular spines few, averaging 16 (4-41)

See rete ie <a RM TES ce Ts ONT ie a Riayat P. fallisi, n. sp. 10. Ventral band ratio more than 0.80 .............. 11 -- Ventral band ratio less than 0.75 ............... 12

11. Prothoracic fringe averages 150 (80-200) wu; circumstigmatal foidsprrregulariceiy iy caceng ) Bike a kes P. sialia, n. sp.

Protocalliphora

Prothoracic fringe averages 58 (34-90) wu; circumstigmatal foldsiconcentric) 2.94 3 P. spatulata, n. sp.

. Dorsal cuticular ridges pronounced; dorsal cuticular spines

both anteriorly and posteriorly directed ....... 13 Dorsal cuticular ridges weak, dorsal cuticular spines directed posteriorly Only. vie 6.2 ee hee ee 16

. Mesostigmatal area with folds and plaques reduced or absent;

mesostigmatal area entire ..............000% 14 Mesostigmatal area with pronounced folds and/or plaques bisecting the stigmatal area ................ 15

. Prothoracic fringe averages 74 (31-130) mw; spines of dorsal

cuticle and medial band large, averaging 40 (16-64) p; puparia large (13 mm) ............ P. lata, n. sp. Prothoracic fringe averages 57 (21-103) uw; spines of dorsal cuticle and medial band smaller than above, averaging less than 40 pw (21-41 and 16-37, resp.; puparia medium-sized CDSS. Oem) Pec eke en dane P. hirundo, S. & D.

. Circumstigmatal folds pronounced, irregular .........

RNa LE TD OC ae P. avium, S. & D. Circumstigmatal folds weak, concentric ........... URE Meee ha cies ken Rite ane at Ln P. chrysorrhoea (Meigen)

. Mesostigmatal folds and/or plaques bisecting stigmatal plate

ri dor: bagel He en ia AIS tS Mie een Noack oy Rina oral 17 Mesostigmatal folds and/or plaques weak or absent, not markedly bisecting stigmatal plate area ........ 19

. Prothoracic fringe averages 75 (55-110) ; ventral band ratio

OPS LE eH OO Ie We Reis P.