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. hesperioides, n. sp.
Prothoracic fringe averages less than 55 ; ventral band ratio
OSMOWOR TESS iS 0ey. Sova es ee lionel Guta een ate: aereaerac eye 18

. Circumstigmatal folds pronounced; dorsal cuticular ridges

ADSEN Ey SRO ie P. seminuda, Nn. sp.
Circumstigmatal folds weak or absent; dorsal cuticular ridges
weak, but always present ....... P. spenceri, n. sp.

. Hyperstigmatal spines small, averaging 11 (4-24) uw;

hypostigmatal spines as tubercles, appearing as fine
SANG PAPER fe) ai Fiaye ls cal a ee eae P. shannoni, n. sp.

P.aenea 93

-- Hyperstigmatal spines large, averaging over 25 U;
hypostigmatal spines as spines .............. 20

20. Mesostigmatal folds and plaques reduced; posterior band
reduced but present; prothoracic fringe averages 60
CZOSSO) ARTES LET IIT CRE NE P. cuprina (Hall)

-- Mesostigmatal folds and plaques pronounced; posterior band
pronounced on all segments; prothoracic fringe averaging
MIOTERtMANN TS Ll A I es er eric.) 2S 21

21. Ventral band ratio less than 0.60 ..... P. bicolor, n. sp.
-- Ventral band ratio more than 0.70... P. parorum, n. sp.

Protocalliphora (P.) aenea Shannon and Dobroscky
(Figs. 12, 42, Map 4)

Protocalliphora splendida var. aenea Shannon and Dobroscky,
1924: 250, 251 [N.H., Ont., latter now known to be braueri;
the N.H. specimens were mentioned in Townsend, 1919: 379
as atypical metallica].

P. splendida form aenea.--Johnson, 1925a: 216 [N.H. record
repeated].

Apaulina aenea.--Hall, 1948: 185 [Description mixed with hirudo
(i.e., braueri); B.C. and Mont. specimens are braueri, and
description of male is of braueri].

Protocalliphora aenea.--James, 1955: 24 [No Calif. specimens
seen, though listed by Hall (1948) from central Calif.]; Boyes,
1961: 552, 554, 556, 565 [Cytology]; Boyes and van Brink,
1965: 543 [Cytology]; Hall, 1965: 925 [Catalog; widespread];
Boyes and Shewell, 1975: 446 [Mention of cytology];
Whitworth, 1976: 17, 26, 37 [Utah; parasite of American
dipper].

Diagnosis. Conspicuously dichromatic, thorax and abdomen
shining dark blue to bluish purple in male, aeneous or dull
cupreous green in female; superficially similar to braueri but
without accessory notopleural bristles, with reclinate orbital
bristles in female, and without the distinctive male terminalia of
braueri.

Male. Head with parafrontal and parafacial dark gray to
brownish gray microtomentose, parafrontal darkening toward
vertex; occiput with two to three irregular rows of long black
hairs behind the postocular setae. Body shining dark blue to

94 _ Protocalliphora

bluish purple, scutum and abdomen concolorous, with former
appearing slightly darker from the gray to brownish gray
microtomentum that demarcates three broad shining stripes.
Abdomen highly shining in direct view but with a thin gray
"bloom" when viewed from behind at a low angle. Calypteres
brownish yellow with brown rims, upper calypter slightly paler.

Frons narrow, narrowing slightly anterior to ocellar tubercle,
at narrowest 0.085 times head width (0.07-0.10) and slightly
broader (1.32x; 1.00-1.50) than ocellar span; ocellar tubercle
ending acutely just anterior to median ocellus, usually by less
than length of latter, the short triangular preocellar area polished;
parafrontals narrow and well separated throughout, the frontal
vitta over twice width of a parafrontal, each parafrontal strongly
narrowing above and becoming subshining, the microtomentum
thin and the prevertical area partly polished; upper portion of
parafacial and the adjacent parafrontal with numerous coarse
black hairs, on the parafacial in three to four irregular rows; each
parafacial of moderate width, 1.33 times (1.18-1.67) width of
frons, 1.75 times (1.50-2.02) ocellar span, and 0.70 times
(0.64-0.83/24) vibrissal interval; cheek height 0.40 times
(0.37-0.42/24) maximum length of eye.

Thorax without accessory notopleural bristles; tympanic
membrane typically bare or with only one or few slender and
rather inconspicuous black hairs, never densely beset with coarse
hairs; postalar wall typically bare, sometimes with a few short
hairs. Fore tibia with one median posterior bristle, rarely a
second.

Terminalia not distinctive, of a type common to a number
of species: cerci distally acuminate and parallel (as in Fig. 5b);
surstylus in profile digitate, slightly broadened toward base but
distally with sides subparallel and apex bluntly rounded (Fig.
12); distal portion of aedeagus slender, not cylindrical as in
braueri (cf. Figs. 6a,c).

Female. Body aeneous or dull cupreous green, rather heavily
microtomentose, dorsum of thorax more heavily so than abdomen
but scutum with three subshining stripes, the abdomen often
quite shining aeneous to cupreous. Calypteres usually yellowish
tinted, rims strongly so.

Frons broader than in braueri, at vertex over 1/4 width of
head (0.26x; 0.24-0.295) and 3.41 times (2.86-3.91) ocellar span;
frontal vitta with a few weak and inconspicuous hairs, heavily
microtomentose, often with small, polished, irregular to
triangular preocellar area; parafrontal relatively broad, with one
to two irreguiar rows of hairs, entirely microtomentose

P.aenea 95

throughout, the prevertical area darker but still evenly
microtomentose; parafacial well covered with hairs, width 0.48
times (0.45-0.55) width of frons, 1.64 times (1.43-2.00) ocellar
span, and 0.78 times (0.67-0.89/23) vibrissal interval; cheek
comparatively narrow, 0.40 times (0.37-0.48) maximum length
of eye.

Apparent normal length of adults, 7.5-9 mm.

Egg. Length 1130 wu (1080-1190)/18, breadth 320 pw
(295-380)/18; cuticle white in color and ornamented with minute
spines.

Second instar. Cephalopharyngeal skeleton 510 Uw
(500-524)/2 in length. Prothoracic fringe 35 b (26-54)/30.
Posterior region: peritreme closed, stigmatal plates 77 wu
(69-86)/4 in diameter; button indistinct; mesostigmatal spines
absent; hyperstigmatal spines numerous, 10 pm (4-14)/30;
hypostigmatal spines scarce, 6 (2-10)/30. Dorsal cuticle with
numerous spines, 12 mw (6-18)/30. Ventral cuticle: spines of
anterior patch 5 pu (4-6)/30; medial band pronounced, spines 8
(4-12)/30; posterior band present.

Third instar. Peritreme usually closed but sometimes open;
mesostigmatal and hypostigmatal spines present. Prothoracic
fringe 78 pw (50-120)/100. Cephalopharyngeal skeleton 805 pw
(740-1010)/100 in length.

Puparium. Length 8.1 mm (5.7-9.7)/100, breadth 3.8 mm
(2.2-4.7)/100; thin-walled, dull, light brown. Posterior region
(Fig. 42a,b): stigmatal plates 170 wu (130-220)/192 in diameter;
distance between buttons 545 pw (340-690)/96, and across
stigmatal plates 1100 w (780-1310)/96; stigmatal ratio 0.49
(0.43-0.52)/96; stigmatal area weakly outlined in cuticular
plaques; mesostigmatal spines present, mesostigmatal folds and
plaques absent; hyperstigmatal spines 34 wm (16-52)/100;
hypostigmatal area with vestigial cuticular folds and numerous
true spines, 15 wu (6-27)/100; circumstigmatal folds absent.
Dorsal cuticle (Fig. 42d) with numerous spines, 24 pu (14-41)/100;
cuticular ridges weak or absent. Ventral cuticle (Fig. 42c): ventral
band ratio 0.47(0.45-50)/100; spines of anterior patch 13 pu
(6-23)/100; medial band present, spines 21 pw (8-33)/100;
posterior band vestigial or absent (if present, as a few rows of
spines in a short arc about the scar of the succeeding ventral
pad).

Type Series. Holotype and one paratype, Franconia, N.H.
(Mrs. Slosson), and one paratype, "N.H.," all females [USNM].
The original description (p. 251) was vague on the type series,
saying only "Type.--Cat. No. 26860 USNM." Considering the

96 Protocalliphora

entire text, however, aenea is listed in the key as from N.H. and
Ont., and on p. 252 the authors spell out the details of three
females from Franconia, N.H. (Mrs. Annie T. Slosson) and one,
probably female, fragmentary specimen from Ontario (And.
Halket). For those who would not accept the "Type No." as a
designation of holotype, Hall (1948) gave the data of the
specimen in full and called it "Type", and that would serve as
lectotype designation under Article 74. The broken specimen
from Ontario was not labeled paratype by Shannon and
Dobroscky and probably was not considered as such because of
its condition; it is after all a female braueri (cf. succeeding
discussion and that under braueri).

Specimens Examined. 202 males, 222 females; major sources,
Gothic, Colo., 89 males, 110 females, and Algonquin Park, Ont.,
94 males, 79 females.

Distribution. Northern, transcontinental, but not often
recorded: Canada and northern United States, south in the
mountains to California, Colorado, and Virginia. Recorded by
Hall (1948) "from central California," but James (1955) recorded
none in his study of "The blowflies of California," and we have
seen only a recently collected series from Fresno County,
California.

Canada. British Columbia: Bowser; Kleanza Creek (14 mi E
Terrace); Shames (16 mi SW Terrace); 32 mi SW Terrace;
Whytecliff (W side of West Vancouver). Ontario: Algonquin Park
(chiefly Lake of Two Rivers) (mostly from nests of barn swallow,
also of common grackle, eastern phoebe, American robin, house
wren). Quebec: Burn Creek near Baie Comeau; Duchesnay (25 mi
W of Quebec City); Hull; Sept. Iles; Val d’Espoir; Sully; Duncan
Lake near Rupert (phoebe).

Contiguous United States. California: Near Bear Creek in
Fresno Co. (nest of American dipper). Colorado: Gothic
(Gunnison Co., N of Crested Butte, 9500 ft, nest of American
dipper; Idaho Springs (6 mi SW, 9000 ft); Weld Co. (nest of Say’s
phoebe); SW corner Jackson Co., 10 mi N junction of highways
Colo. 14 and US 40. Idaho: Wallace. Maine: Little Black River
Rapids (near mouth of Allegash River), published as P. splendida
splendida by Johnson (1925a:6). New Hampshire: Franconia
(including type series); Mt. Washington. New York: Whiteface
Mt. in Essex Co. Oregon: Mary’s Peak (W of Corvallis; paratype
of cuprina Hall). Utah: Temple Fork in Cache Co. near Logan
(nest of American dipper). Virginia: Shenandoah National Forest
(nest of phoebe; puparia only); White Oak Canyon in Shenandoah
National Park (nest of Louisiana waterthrush). Washington: Deer

P.aeneaa 97

Lake near Chevelak. West Virginia: 12 mi SW of Berkeley
Springs (nest of phoebe).

Of the hand-collected material, the earliest recorded date is
May 14 (Whytecliff, B.C.) and the latest September 8 (Little
Black River, Maine).

Basis of description of immature stages (all material
Algonquin Park, Ontario): Egg: Algonquin Park, Ont., from
caged female, July 1955. Larvae: 2nd instar (2 specimens): barn
swallow, June, 1951; 3rd instar (10 specimens): barn swallow,
June, 1951 (7) and eastern phoebe, June, 1954 (3). Puparium
(100 specimens): barn swallow, June 1951 (34); June, 1952 (6);
July, 1954 (20); June, 1955 (10); eastern phoebe, July, 1953 (10)
and July, 1954 (20).

Hosts. American dipper (formerly water ouzel), common
grackle, eastern and Say’s phoebes, American robin, barn
swallow, Louisiana waterthrush, and house wren.

Ecology and Biology. Apparently rare except in certain
limited areas. It has been found frequently in nests at any height
in or around man-made structures, the most frequent hosts being
barn swallows and phoebes. Especially noteworthy are the three
records from the American dipper. The record from Gothic,
Colo. is based on an unusually long series of 89 males, 110
females reared July 15 and 18, 1960 (C. L. Remington), from
"larvae in current nest of Cinclus mexicanus" [Yale]. No other
species of Protocalliphora is known from that species of bird at
the present time. Both ecology and biology of aenea were studied
extensively by Bennett at Algonquin Park.

Variation. In the series of 25 measured females from which
the proportions were derived, there were 18 eastern (chiefly Ont.
and Va.) and only 7 western (B.C. to Colo.). Proportions were
calculated on these separately to determine if the eastern and
western populations showed differences, but differences were
slight. Subsequently a good series from Gothic, Colo., was
received, and the 10 females were measured and proportions
calculated as a test of both the proportions based on the whole
series and of those based on the few western specimens alone.
The comparative proportions are as follows (Table VII) with their
ranges (all based on numbers of females shown, except for the
parafacial/vibrissal interval, for which two western individuals
could not be measured. While the means differ, in general the
ranges overlap and it appears that there is only one widely
distributed species.

Protocalliphora

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P.asiovora 99

Remarks. Confusion of the aeneous females of aenea and
braueri has apparently been frequent and has led to erroneous
records. The broken specimen of "aenea" from Ontario, recorded
by Shannon and Dobroscky (1924) as if part of the type series
although not labeled a paratype by them, is actually a female
braueri, as shown by the accessory notopleural bristles and the
distinctive puparium, fortunately preserved with the specimen.
The endoparasitic habit of the larva also indicated braueri. The
British Columbia and Montana specimens recorded as aenea by
Hall (1948: 186) were also found to be braueri as noted under that
species. Hall’s description of the male of aenea applies to
braueri, and was probably based, in part at least, on three males
from the Hamilton, Mont. series; the other material cited consists
only of females. The proportions given for the female of "aenea"
were apparently influenced by the series of braueri, and cannot
be relied on for aenea.

Males of aenea have no striking feature or distinctive
combination of characters and will be difficult to identify by
themselves. They have probably often been misidentified in
collections. Females are striking because of the entirely aeneous
coloration, shared only with the female of braueri, from which
aenea is easily separated by having reclinate orbital bristles and
broader front, and no accessory notopleural bristles. Associated
puparia also will at once distinguish braueri and aenea.

Teneral females sometimes lack the distinctive aeneous or
cupreous green color, and tend to a greenish color tinted with
copper. To the naked eye, however, when a series containing
both sexes is examined, the females, even though not fully
colored, are so distinctly different from the males that there is no
question but what a dichromatic species is involved.

Puparia of aenea are readily confused with those of fallisi
and tundrae and its close associates. Those of aenea are generally
distinguished by (i) the long prothoracic fringe, (ii) absence of
mesostigmatal folds, and (iii) occurrence in nests of birds in
man-made environments.

Protocalliphora (P.) asiovora Shannon and Dobroscky
(Figs. 8, 35, 43, Map 5)

Protocalliphora avium var. asiovora Shannon and Dobroscky,
1924: 249, 250 [Wash.]; Shannon, 1926: 128 [In key].

P. avium.--Jellison and Philip, 1933: 28-30 [Mont.]; Gahan, 1933:
31-33 [New chalcidoid parasite].

100 = Protocalliphora

New genus and n. sp.--Neff, 1945: 73-76 [Calif.].

Apaulina avium in part.--Hall, 1948: 186 [asiovora in synonymy
under avium].

Apaulina basingeri Hall, 1948: 190 [Calif; n. sp. of Neff 1945.].
Protocalliphora \arvae.--Hill and Work, 1947: 74-75 [Calif., on
sparrow hawk [i.e., American kestrel] and golden eagle].

P. avium.--Pletsch, 1948: 296-7 [Mont.]; Jellison, 1969: 55
[Mont.].

P. sp.--Lee and Ryckman, 1954: 24 [Ariz.].

P. avium and basingeri.--James, 1955: 25 [Calif. records all under
avium (one from Big Dalton Dam is correctly hirundo);
basingeri probably a synonym of avium].

P. avium.--Gill, 1956: 652 [Alaska]; Kamal, 1958: 262 [Idaho].

Calliphora sp. in heads of prairie falcons.--White, 1963: 442-3
[Utah; probably P. asiovora].

Protocalliphora asiovora.--Hall, 1965: 926 [B.C. to Mont., s. to
Calif., N. Mex., and Colo.; basingeri a synonym teste
Sabrosky and Bennett].

P. asiovora.--James, 1969: 252 [Mention of biology].

P. sp. on hawks.--Seidensticker and Reynolds, 1971: 411 [Mont.;
probably asiovora].

P. asiovora.--Whitworth, 1976: 22ff [Utah]; Whitworth, 1977:
4933-B.

Diagnosis. As in P. avium but frons of male slightly
narrower; western, sister species of avium.

Male. Color as described for avium. Frons (Fig. 35)
moderately broad at vertex, but slightly narrower than in avium,
0.13 times head width (0.115-0.14; holotype of basingeri 0.14)
and much less than twice ocellar span (1.68x; 1.44-1.88);
preocellar area often granular or weakly shining; parafacial
broad, as in avium, slightly wider than frons (1.14x; 1.00-1.31),
obviously much wider than ocellar span (1.93x; 1.75-2.20), and
usually slightly wider than vibrissal interval (1.04x; 0.91-1.14);
cheek broad, its height nearly half maximum length of eye
(0.47x; 0.44-0.52).

Thorax without accessory notopleural bristles and hairs on
tympanic membrane; postalar wall bare. Fore tibia with one
posterior bristle.

Terminalia: cerci sturdy, deeply notched, widely divergent in
posterior aspect (cf. Fig. 5); surstylus broadly rounded, similar to
avium but slightly broader in proportion to its length (Fig. 8);
aedeagus similar to the of avium (cf. Fig. 6a).

P.asiovora 101

Female. Color as in avium; concolorous with male but
dorsum of thorax more heavily gray microtomentose.
Microtomentum of parafacial and parafrontal brownish yellow,
not silvery as in male; prevertical area of parafrontal dull, dark
gray microtomentose; frons at vertex moderately broad, 0.295
times head width (0.28-0.32) and 3.52 times ocellar span
(3.00-4.29); frontal vitta often with some strong bristlelike setae;
preocellar area dull; parafacial broad, its width slightly over half
that of frons (0.56x; 0.50-0.59), 1.96 times ocellar span
(1.75-2.27), and 1.13 times vibrissal interval (1.00-1.24/24);
cheek height nearly half maximum length of eye (0.49x;
0.42-0.525).

Apparent normal length of adults 8.5-9 mm, commonly about
9. A very few specimens, no doubt underfed, are as short as
6.5-7 mm.

Third instar. Peritreme open. Prothoracic fringe rather
sparse, 32 st (8-66)/72. Cephalopharyngeal skeleton 980 wu
(850-1080)/11 in length.

Puparium. Length 8.9 mm (8.6-9.1)/6, breadth 4.1 mm
(3.6-4.4)/13; medium-thick walled, dull, dark brown. Posterior
region (Fig. 43a): stigmatal plates 200 mw (150-220)/26 in
diameter; distance between buttons 500 pm (460-590)/13, and
across stigmatal plates 1200 uw (1000-1390)/13; stigmatal ratio
0.40 (0.36-0.50)/13; stigmatal area not prominently outlined in
cuticular plaques; mesostigmatal spines present; mesostigmatal
area without plaques but with folds prominently bisecting
stigmatal region; hyperstigmatal spines 24 wm (8-31)/150;
hypostigmatal area with numerous, prominent cuticular folds;
hypostigmatal spines 15 fw (6-27)/150; circumstigmatal folds
present, weak and concentric (in some cases, barely discernible
and considered absent). Dorsal cuticle (Fig. 43b) with numerous
spines, 25 wb (12-45)/150; cuticular ridges weak. Ventral cuticle
(Fig. 43c, d): ventral band ratio 0.63 (0.50-0.69)/10; spines of
anterior patch 14 w (8-27)/150; medial band pronounced, but
interrupted on midline by pronounced break, spines 20 wu
(10-31)/150; posterior band pronounced.

Types. Type series of asiovora: Holotype male, McElroy
Lake, Paha [Adams Co.], Wash. June 30, 1920 (R. C. Shannon),
reared from larva found in nest of long-eared owl. Type No.
26858 in the U.S. National Museum of Natural History.

Type series of basingeri: Holotype male, San Diego, Calif.,
June 28, 1917 (J. M. Aldrich). Type No. 54929 in the USS.
National Museum of Natural History. Paratypes (all California):
2 males, same data as holotype [USNM]; 4 males, Temecula,

102 Protocalliphora

Riverside Co., June 4, 1936 (A. J. Basinger), one bearing a
penciled note "from sunshine on bridge" [USNM]; 2 males, Forest
Home, San Bernardino Co., May 18, 1934 (A. J. Basinger)
[USNM}]: 3 males, 2 females (not 2 males, 3 females as published),
Le Grand, Merced Co., June 10, 1938, from nest of mourning
dove (J. A. Neff) [males, USNM; females, CAS]: 2 females,
Anaheim, Orange Co., July 1936, from nest of house finch (A. J.
Basinger) [USNM, CAS]; male (now headless), Corona, Riverside
Co., Sept. 20, 1934 (Basinger) [CAS]; 6 males, San Diego, Sargent
(Santa Clara Co.), Lockwood (Monterey Co.), and Laguna (San
Diego Co.) (R. H. and Jack Beamer)[Kans. U.]. Published
paratype not found: female, Riverside, Calif. (Basinger).

Specimens Examined. 315 males, 255 females.

Distribution. British Columbia to Montana, south to Baja
California and New Mexico. This species should also occur down
the Mexican Plateau.

Canada. British Columbia: 2 males, 2 females, Kamloops
(American robin nest), Laggan (old railroad station for Lake
Louise), Oliver, and Orofino Mt. (4000 ft) near Oliver.

Contiguous United States. Arizona: 17 males, 8 females,
Chiricahua Mts. (8000 and 8200 ft) near Portal, Coconino Co.
near Flagstaff (nest of great horned owl), Globe, Huachuca Mts.
(Ramsey Canyon), Doney Park near Flagstaff (K. A. Walker; total
of 55 larvae removed from "various external portions of a dead
nestling" as well as from nest material). California (by county):
48 males, 32 females, Alameda, Contra Costa, El Dorado, Kern,
Los Angeles, Madera, Merced, Modoc, Mono, Monterey, Orange,
Riverside, San Bernardino, San Diego, Santa Clara, Shasta,
Tulare, and Tuolumne Counties, from nests of mourning dove
(Le Grand in Merced Co.), house finch (Anaheim in Orange Co.),
great horned owl (O’Neals in Madera Co.), and barn swallow
(Adin Bridge in Modoc Co.). Also larvae from Alameda Co.
(Sunol, golden eagle), and Santa Clara Co. (Milpitas, two removed
from flesh at bottom of crop of golden eagle, one from under left
wing of golden eagle, and 5 from ear of great horned owl). Also
puparia, Merced Co. (Le Grand, from nest of loggerhead shrike).
Colorado: 12 males, 6 females, 16 larvae, Boulder, Colorado
Springs (black-billed magpie and horned owl nests), Platte
Canyon (near Idlewood). Idaho: 6 males, 4 females, Franklin in
Franklin Co., Lake Waha, Moscow Mt. Also larvae from Boise,
Elmore, and Owyhee Counties (all in golden eagle nests). Also
larvae on black-billed magpies, Moscow, published by Kamal
(1958), probably asiovora. Montana: 185 males, 185 females,
Connor, Hamilton (European starling nest), Gallatin Co. (Fairy

P. asiovora 103

Lake, 9500 ft), Beaverhead Co. (Dillon, long-eared owl nest),
Beaverhead Co. (nests of American crow, black-billed magpie,
long-eared owl, and Swainson’s hawk), Granite Co. (swallow
nest), and Ravalli Co. (American crow and black-billed magpie
nests; published Jellison and Philip 1933). Nebraska: male,
Dawes Co., 8-11 mi SE Crawford [See note at end of Remarks].
Nevada: male, Douglas Co. (Dagget Pass). New Mexico: 3 males,
Bernalillo (Cedro Canyon), Otero (Weed), and Torrance (Tajique)
Counties. Oregon: 6 males, Dufur, Corvallis (cliff swallow nest).
Camp Abbot in Deschutes Co., 7 mi SW Beatty in Klamath Co.
Utah: 28 males, 12 females, Brigham City (Bear River Refuge,
black-billed magpie nests), Box Elder Co. (Malaise trap), Cache
Co. (Benson, nest of Brewer’s blackbird, chiefly infested by
metallica), Salt Lake City (black-billed magpie nest), and
Wasatch Mts. Washington: 5 males, 6 females, Almota (Shannon
and Dobroscky 1924), Central Ferry, McElroy Lake at Paha (nest
of long-eared owl, holotype of asiovora), Okanogon Co.
(black-billed magpie nest), Pullman, Wenatchee. Wyoming:
female, South Fork of Powder River, 1.8 mi W of town of
Powder River in Natrona Co.

Mexico: male, Baja Calif., Sierra San Pedro Martir, La Grulla,
6900 ft, June 15, 1953 (P. H. Arnaud, Jr.) [CAS].

Basis of description of immature Stages: Larva, 3rd instar
(5 specimens): Ravalli Co., Mont., black-billed magpie, May 28,
1931. Puparium (13 specimens): Le Grand, Calif., mourning
dove, June, 1939 (4) and 1943 (6), loggerhead shrike, 1943 (1);
Okanogon Co., Wash., black-billed magpie, June, 1939 (2).

Hosts. Brewer’s blackbird, American crow, mourning dove,
golden eagle, house finch, several hawks (northern goshawk,
Cooper’s, ferruginous, red-tailed, and Swainson’s), pinyon jay,
black-billed magpie, great horned and long-eared owls, common
raven, American robin, loggerhead shrike, European starling, and
barn and cliff swallows. Probable host: American kestrel
(sparrow hawk), published by Hill and Work (1947).

Ecology and Biology. The host records thus far known
suggest that asiovora generally occurs at intermediate or forest
canopy levels and usually infests large birds and often raptors,
but it occurs more often in smaller birds than does its sister
species in the East, P. avium.

Variation. Typically, asiovora has only one posterior bristle
on the fore tibia, but occasional specimens have two, usually on
one side only, rarely two on both sides. In a series of 82 from
three nests in Ravalli Co., Mont., seven specimens had extra

104 = Protocalliphora

bristles. Six of these had two on one side and one on the other,
and one had two on each fore tibia.

Remarks. P. asiovora is a sister species and western
equivalent of the eastern P. avium. The two are very similar in
general appearance and in the broad parafacials and exceptionally
broad surstylus of the males. Females have proved impossible to
distinguish except by their occurrence with males and within the
characteristic ranges for the two species. The character, shared
only with interrupta, of an interrupted median ventral spine band
in the larvae, supported by the consistently narrower frons in the
male asiovora and the different distributions of the two, have led
us to recognize them as distinct species.

Protocalliphora asiovora is one of only three species with
broadly rounded surstylus in the male (Figs. 7-9), the others
being the sister species, avium, and P. deceptor. P. avium and
asiovora are very close, and the latter was originally described as
a mere variety of avium. The sharpest distinction here, and the
main reason why we recognize asiovora as a distinct species, is
that the medial spine band on the ventral surface of mature
larvae (and of course, puparia) is distinctly interrupted on the
midline (Fig. 43c,d). This spine band is continuous in avium.
Both avium and asiovora have the mesonotum in both sexes more
heavily microtomentose than in most other species, especially in
the males. One finds it more appropriate here to speak of two
broad gray stripes that set off median and lateral shining stripes,
rather than to speak of three shining stripes.

The third species with broadened surstylus, P. deceptor, is
quite a different species, with narrow frons in both sexes, as
narrow as the frons in P. braueri, and narrow parafacial. It is a
species of eastern United States, and chiefly parasitizes small
passeriform birds. It is also worth noting that the cerci in
deceptor are narrow, acuminate, and parallel to their apices, quite
unlike the sturdy and divergent cerci of avium and asiovora (cf.
Figs. 5a and Sb).

Larvae and puparia of asiovora are recognized from all other
Protocalliphora except interrupta by the marked interruption of
the medial band (Fig. 43c, d). They are separated from interrupta
by (i) the larger ventral band ratio (0.63, cf. 0.27 for interrupta),
(i1) the larger spines on the cuticle which appear as spines, not as
small tubercles as in interrupta, and (iii) the pronounced
mesostigmatal folds in asiovora compared to their absence in the
latter species.

It is probable that asiovora is sometimes involved in myiasis
like its close relative, the eastern P. avium, for which there are

P.avun 105

many records of aural myiasis in particular. This may be
principally the result of greater attention to nestling raptors in
eastern North America. Specimens of asiovora from Milpitas,
Calif. were taken from golden eagles from flesh near the bottom
of the crop, and from under the left wing, also from the ear of
a great horned owl. Hill and Work (1947) in California reported
Protocalliphora larvae in the ears and nostrils of sparrow hawk
(i.e., American kestrel) and golden eagle, and from this larval
habit we can be reasonably sure that they were asiovora. The ear
canal of the hawk was swollen and filled with squirming larvae,
and the nostrils were deformed. In Utah, Whitworth often
observed asiovora in aural myiasis.

Two males from the northwestern corner of Nebraska are
somewhat intermediate between asiovora and avium. In the one
from Dawes County the frons appears narrow as in asiovora, the
ratio of frons to head width being 0.14, at the upper end of the
range for that species. The surstylus and cerci are like those of
asiovora. In the one from Sioux County, the ratio of frons to
head width is 0.16, which would agree with that of avium. The
male terminalia are unfortunately lacking in this specimen. It
would be interesting to have reared series, with immature stages
preserved, to solve questions of identity: Do the two species
actually overlap here? Or are these specimens extreme variants
of one or the other? For the present, each is recorded under
what appears to be its species, but with attention called to the
need for further study and the possibility of mistaken
identification.

Protocalliphora (P.) avium Shannon and Dobroscky
(Figs. 5a, 6a, 7, 34, 44, Map 5)

Protocalliphora azurea (Fallén).--Coutant, 1915: 135-150 [N.Y..,
on crow].

Larvae "possibly" Compsomyia macellaria (Fabricius).--Burtch,
1915: 293 [N.Y., in ears of Cooper’s hawk; undoubtedly P.
avium].

Protocalliphora avium Shannon and Dobroscky, 1924: 250 [N.Y..,
from crows].

P. avium.--Dobroscky, 1925: 275 [N.Y.]; Shannon, 1926: 128 [in
key]; Johannsen, 1928: 827 [N.Y., Coutant material].

P. sp., larvae in ears of hawks.--Bent, 1937: 189-190 [Probably
N.H.; undoubtedly P. avium].

106 =‘ Protocalliphora

P. splendida sialis [sic].--Sargent, 1938: 82-4 [N.Y., nests of
hawks, some larvae in ears; undoubtedly P. avium].

P. sp., larvae on peregrine falcon.--Hickey, 1942: 189 [N.Y.,
probably P. avium}.

Apaulina avium.--Hall, 1948: 186-190 [N. U.S. and s. Canada;
asiovora here a synonym].

Protocalliphoria [sic] larvae, ears of hawks.--Hamerstrom and
Hamerstrom, 1954: 4-8 [Iowa, Wis., Mich.; undoubtedly P.
avium].

Apaulina avium.--Meng, 1954: 21-22 [N.Y., biology].

Protocalliphora avium.--Boyes, 1961: 552ff. [Cytology]; Boyes
and van Brink, 1965: 543 [Cytology]; Hall, 1965: 926 [Wis.,
Ont., N.Y., Pa.].

P. sp., larvae in ears of young peregrine falcons.--Hagar, 1969:
127 [Mass.] and Hickey, 1969; pl. 43 [Vt.] [Both undoubtedly
P. avium).

P. avium.--Boyes and Shewell, 1975: 446 [Mention of cytology];
Bohm, 1978: 297 [Minn., in ears of great horned owls];
Tirrell, 1978: 21-5 [N. Dak., on hawks and owls].

P. avium.--Bortolotti, 1985: 165-8 [Frequency on bald eagles,
Sask.].

P. avium.--Poole and Bromley, 1988: 36 [On gyrfalcon nestling,
Central Canadian Arctic].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic dark blue to bluish purple; calypteres white; frons of
male of moderate width; parafacial broad in both sexes; surstylus
of male broadly rounded (Fig. 7); eastern, sister species of P.
asiovora.

Male. Head chiefly black in ground color, frontal vitta and
genal groove reddish, and parafrontal and parafacial bright gray,
silvery microtomentose, that of parafrontal posteriorly thinning
to shining black opposite ocellar tubercle, parafacial without
changeable reflecting spot. Body shining metallic blue to bluish
purple, more or less gray microtomentose, heavily so on scutum
and scutellum, especially in two broad stripes (as viewed from
behind) that set off a median shining stripe and the lateral
shining areas, each of which may be weakly separated into a
lateral stripe and a supraalar vittula; abdomen appearing shining
in direct view but 2nd to 4th tergites gray microtomentose when
viewed from behind at a low angle, Sth tergite always shining
and non-microtomentose. Calypteres opaque whitish, outer rims
pale yellowish.

P.avium 107

Frons moderately broad (Fig. 34), at vertex 0.16 times head
width (0.15-0.18), and nearly twice ocellar span (1.96x;
1.68-2.19/25); frontal vitta broad throughout, at narrowest twice
or more width of a parafrontal; preocellar area granular and
subshining or weakly shining; parafrontal distinct throughout, the
narrowed upper half with distinct row of fine hairs outside
frontal bristles, the row becoming 2-3 irregular rows anteriorly
and merging with the numerous long black parafacial hairs;
parafacial notably broadened for Protocalliphora, approximately
equal to width of frons at vertex (0.955x; 0.86-1.06), obviously
much wider than ocellar span (1.87x; range 1.56-2.27), and
slightly wider than vibrissal interval (1.16x; range 1.00-1.29);
cheek broad, nearly half maximum length of eye (0.45x; range
0.42-0.48).

Thorax without accessory notopleural bristles and without
hairs on tympanic membrane; postalar wall usually bare,
sometimes with a few black hairs. Fore tibia usually with one
posterior bristle.

Terminalia: cerci distally slender and tapering, divergent in
posterior view (Fig. 5a), strongly curved in profile (Fig. 7);
surstylus broadly rounded, subovate (Fig. 7); aedeagus as usual
for the genus (Fig. 6a).

Female. Color as described for male, but both thorax and
abdomen more heavily gray microtomentose, especially the
former. Microtomentum of parafrontal and parafacial dark gray
to brownish yellow, not silvery as in male; preocellar area dull,
densely brownish microtomentose; prevertical area of each
parafrontal dull, dark gray microtomentose; frons at vertex
moderately broad, 0.295 times (0.27-0.32) head width and 3.45
times (2.89-3.88) ocellar span; frontal vitta usually with a few
strong black mesoclinate setae in addition to weaker and
inconspicuous hairs; parafacial unusually broad for the genus,
width slightly over half that of frons at vertex (0.54x; 0.50-0.59),
1.86 times (1.56-2.25) ocellar span, 1.15 times (1.00-1.36/21)
vibrissal interval, and about twice width of 3rd antennal segment,
usually slightly convex and appearing bulging; cheek nearly half
maximum length of eye (0.47x; 0.42-0.50).

Apparent normal length of adults, 9.5-10 mm.

Egg. Length 1040 w (950-1150)/33; breadth 340 wu
(275-405)/33; cuticle of yellowish color; cuticle with spinules.

Second instar. Cephalopharyngeal_ skeleton 630 [UL
(520-730)/5 in length. Prothoracic fringe 15 s (10-26)/50.
Posterior region: peritreme closed, stigmal plates 94 wp
(86-135)/10 in diameter; button indistinct; mesostigmatal spines

108 Protocalliphora

scarce or absent; hyperstigmatal spines numerous, 12 pt (6-21)/50;
hypostigmatal spines numerous, 6 (4-12)/50. Dorsal cuticle
with numerous spines, 10 (6-21)/50. Ventral cuticle: spines of
anterior patch 4 pw (2-12)/50; medial band pronounced 8 p
(4-14)/50; posterior band present.

Third instar. Peritreme usually open but sometimes weakly
closed; spines small and scarce between stigmatal plates,
numerous and larger in the hypostigmatal area. Prothoracic
fringe 45 ww (31-61)/100. Cephalopharyngeal skeleton 1080
(870-1260)/72 in length.

Puparium. Length 9.5 mm (6.3-10.5)/100, breadth 3.8 mm
(2.2-4.7)/100; thick-walled, dull, black-brown. Posterior region
(Fig. 44a, b): stigmatal plates 215 w (150-260)/196 in diameter;
distance between buttons 530 mw (325-655)/100 and across
stigmatal plates 1330 mw (860-1620)/100; stigmatal ratio 0.40
(0.38-0.42)/100; stigmatal area outlined in plaques; mesostigmatal
spines scarce or absent, mesostigmatal plaques and folds present;
hyperstigmatal spines 20 pu (10-33)/100; hypostigmatal area with
few, prominent cuticular folds; hypostigmatal spines numerous,
12 mw (6-23)/100; circumstigmatal folds present, pronounced,
irregular. Dorsal cuticule (Fig. 44d) with numerous spines, 23
(10-33)/100; cuticular ridges prominent, usually with two rows
of spines associated with each ridge, one each directed posteriorly
and anteriorly. Ventral cuticle (Fig. 44c): ventral band ratio 0.75
(0.73-0.86); spines of anterior patch 15 ww (6-21)/100; medial
band pronounced, spines 20 pw (10-33)/100; posterior band
pronounced, present on all segments.

Type series. Holotype male, Ithaca, N.Y., reared from
nestling American crow (I.D. Dobroscky), "Cornell U. Lot 1033,
Sub 108A," Type No. 26857 in the U.S. National Museum of
Natural History. Allotype, labeled only "Cornell U. Lot 1033 Sub
108." Paratypes: male, female, same data as type but Sublot 108,
the male labeled May 29, 1923, the female "raised on robins, June
28, 1923" [USNM]; one male, same data as type, and 2 females,
Sublot 108, all labeled March 29, 1923 [Cornell U.]. The original
description recorded three males, five females, but one female
has not been located. The date March 29 was undoubtedly that
of the collection of nest and larvae. The holotype and one male
paratype [Cornell U.] were each pinned with puparium
associated. The holotype bears only the Lot and Sublot label, and
the Type Number label, but the other information was published
with the original description. The Type No. system is by some
not considered designation and it must be admitted that here the
Original description did not specify the sex of the type nor which

P.avum 109

Sublot was involved. To obviate any further argument, I
designate the "holotype" as lectotype.

Specimens Examined. 176 males, 212 females.

Distribution. Northern and northeastern, Alaska and Yukon
Territory, Northwest Territories, Saskatchewan, and North
Dakota to Quebec and Connecticut, south to Nebraska and
Pennsylvania.

Alaska: The material for two identifications of "avium," the
first by M. T. James and the second probably also, could not be
located for verification: Fairbanks (G. Gill), published Gill
(1956); Matanuska (W. J. Chamberlin), in Chamberlin’s
mimeographed report on his Alaska work for 1945.

Canada. Manitoba: 3 females, Bird’s Hill Provincial Park
(from ear of young red-tailed hawk), Churchill ("visiting
Solidago"). Northwest Territories: larvae (det. Bennett),
Mackenzie District, on mainland S of Cambridge Bay and E of
Bathhurst Inlet, 68.1° N and 106° 35’ W (Kim G. Poole) (upper
breast, ears, nostrils, around anus, etc., of gyrfalcon chicks, and
from top of wing, beneath wing, left tarsometatarsus, and around
cloaca of nestling peregrine falcons). Ontario: 57 males, 117
females, larvae and puparia, Algonquin Park (at or near Lake
Sasajewan) (nests of gray catbird, American crow, goshawk,
broad-winged hawk, American robin), Burleigh Falls, Dryden,
Toronto (American crow nest), Osgoode (American crow nest).
Quebec: 2 females, near Eardley, and La Verendrye Provincial
Park. Saskatchewan: male, Besnard Lake at 55° 20’ N, 106° W
(collected as larva in ear of nestling bald eagle). Yukon
Territory: male, Dempster Highway, Mile 87, July 8-12, 1973 (G.
& D. M. Wood) [CNC].

Contiguous United States: Connecticut: male, Stamford.
Illinois: male, female, McHenry Co., from ear of red-tailed
hawk. JIowa: Ruthven, Palo Alto Co., larvae reported by
Hamerstroms (1954). Minnesota: larvae and adults, Trommald
(ghost town near Ironton) (larvae from head and neck region of
bald eagle), near Foley (in ears of great horned owl) (Bohm
1978). Nebraska: male, Monroe Creek Canyon in Sioux Co. NNW
of Harrison [See note at end of Discussion under asiovora]. New
York: 66 males, 55 females, many larvae, Chautauqua Co. (from
ears of broad-winged hawk), Ithaca (American crow nests),
Candor, Catskill Mts., "New Albany" (presumed labeling error for
New Albion in Cattaraugus Co.) (from ear of nestling hawk).
North Dakota: 34 males, 19 females, 38 larvae, 14 localities in
Grand Forks Co. (nests of red-tailed and Swainson’s hawks and
great horned owl). Pennsylvania: 10 males, 11 females,

110 Protocalliphora

Coraopolis (from larvae in ear canals of nestling broad-winged
hawk), State College, Tionesta in Forest Co. Wisconsin: 4 males,
4 females, Monroe in Green Co. (from ear of two-week old
red-tailed hawk), Waushara Co. (from ear of red-tailed hawk),
"Rainbow Flowage" near town of Lake Tomahawk in Oneida Co.
(larvae from ear of osprey).

Basis of description of immature stages: Egg: Algonquin
Park, Ont., from caged female, July 1955. Larvae, 2nd instar (5
specimens): Algonquin Park, Ont., American crow, June, 1954;
3rd instar (10 specimens): Algonquin Park, Ont., American crow,
June, 1955. Puparium (100 specimens): Algonquin Park, Ont.,
American crow, June 1952 (25) and June, 1955 (25),
broad-winged hawk, August, 1955 (25); Toronto, Ont., American
crow, June, 1955 (25).

Hosts. Gray catbird, American crow, bald eagle, gyrfalcon,
peregrine falcon, several hawks (broad-winged, Cooper’s,
red-shouldered, red-tailed, Swainson’s, and northern goshawk
and northern harrier), osprey, great horned owl, and American
robin. The Cooper’s hawk was reported by Burtch (1920) and
Meng (1954), the goshawk and the red-shouldered hawk by Meng
(1954), and the "marsh hawk" (northern harrier) by the
Hamerstroms (1954), all records accepted as probably correct
because of the known characteristic habit of the larvae, and the
eastern localities well apart from asiovora.

The published record of P. avium from swallows in eastern
Canada by Hearle (1938) is uncertain and possibly incorrect. The
wording refers to maggots in cysts on the back of swallows, in
which case it probably refers to braueri. We have no records of
avium from swallows, although there are a few records from
non-raptorial birds.

Ecology and Biology. P. avium is found primarily in the
nests of large birds such as crows and raptors, and usually in
nests in the forest canopy. The larvae are sometimes found
subcutaneously, causing true myiasis, but always in the larger
birds as contrasted with the myiasis-producing larvae of braueri,
chiefly in small birds.

Maggots are often noted in the ears and nostrils of nestling
raptors, as indicated by many of the records, but the exact
damage or extent of damage is seldom recorded. E. C. Schriver
(personal communication) has commented that he had "found
these parasites in the ear openings of all our native tree-nesting
hawks." Burtch (1920) found maggots, probably of avium in the
ears of three nestling Cooper’s hawks. Bent (1937:189-190)
quoted a communication from a correspondent, Lewis O. Shelley,

P.avum = 111

about two young red-shouldered hawks that were taken from a
nest when about three weeks old, and mounted: "When the young
hawks were being skinned both birds were found to have the ears
affected with maggots; 20 were collected from the four ears and
preserved; ... These maggots had eaten about the ears not only to
disfigure the outer ear cavity, leaving the marks of their attacks
so that they are preserved in the mounted specimens, but had
completely destroyed the ear drums of both birds. It is to be
wondered how long these young hawks would have survived had
they lived to reach maturity."

The Hamerstroms (1954) found maggots of Protocalliphora in
the ears of three species of hawks, and they came to suspect that
such maggots "may be a rather common childhood disease of
hawks." In Iowa they found maggots in marsh hawks [i.e.,
northern harriers] "embedded under the wings, near the vent or
on the crown, and especially in the ears and within the nares."
It is noteworthy that the marsh hawk, a ground-nesting species,
represents a radical difference from the usual habit of avium, yet
the larval habit described by the Hamerstroms is
characteristically that of avium. On this basis we accept the
record as probably correct, although we have not seen specimens.
The catbird, not a raptor, is also an unusual host for avium,
nesting as it does at low levels. The nest was three feet above the
ground, and 20 metallica and one avium were reared from it.
Occasional aberrant rearings may occur.

Meng (1954), in an intensive four-year study of Cooper’s
hawks in central New York, found maggots of avium in the ears
of many young birds of this and several other species of hawks.
"When the 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 eyeases are three weeks old their ear openings may
be completely plugged by the protruding ends of the screw-worm
fly larvae. The maggots are intermittent feeders and do not
remain in the ears continuously. At night while the young hawks
are sleeping the hungry larvae find their way into the ears, and
the full ones crawl out into the nest debris. In this way a given
hawk is host to many more parasites then would be indicated by
the dozen or so larvae that are usually found in the ear openings
on an infected bird.". Meng also commented that "None of the
eyeases that had maggots in their ears seemed to be weakened
appreciably, and by the fourth week all the maggots had left the
ears." By that time, of course, the larvae would have matured
and dropped away to pupate.

112 Protocalliphora

Tirrell (1978) in North Dakota found maggots of avium in the
auditory meatus of 34 out of 36 nestling red-tailed hawks, as well
as in one nestling great horned owl and two of Swainson’s hawks.
"Eventually larvae plugged and stretched the auditory meatus.
The ear opening and surface of the ear canal were covered with
a red-brown exudate, and the area surrounding the ear was
inflamed and swollen." Nestlings of the red-tailed hawks were
closely observed, and they were infested between one week and
four to five weeks of age, by which time the avium larvae would
have matured and dropped from the ears.

Bohm (1978) reared avium from larvae in the ear cavities of
nestling great horned owls in Minnesota. Infestations were
recorded on 26 of 46 (54%) of the nestlings banded over a
two-year period. In the same period 41 of 73 (56%) of nestling
red-tailed hawks were also infested, undoubtedly also by avium,
although none were reared and identified.

On a different host, Bortolotti (1985) made repeated
observations (461) individually on 72 bald eagle nestlings in a
three-year period in north central Saskatchewan. He found that
all eaglets had avium larvae "in their ears and on their body at
some time during development." They "frequently were so
numerous that the entire ear opening was blocked. A
conspicuous blackish brown exudate was always present in and
around the outside edge of an infested chick’s ears. Although
this substance became dry, encrusted, and difficult to remove, it
did not persist, for the ears eventually became clean and to.the
naked eye showed no trace of ever being parasitized." Bortolotti
reported that "nestlings were repeatedly parasitized as they grew
older. It was not uncommon to observe a large number of larvae
on a bird during one visit, followed by the absence of larvae on
the next visit a few days later, and then another heavy infestation
on the third visit." The last sentence is especially important in
showing that even the larvae in aural cavities leave the host and
drop into the nest material between feedings. In this study no ill
effects on the birds were observed.

The ecology and biology of avium have been investigated in
some detail by Bennett (1957) at Algonquin Park, Ontario.

Remarks. Protocalliphora avium was adequately diagnosed by
Shannon and Dobroscky and described in considerable detail by
Hall. It is one of the largest species of Nearctic Protocalliphora,
and individual specimens are usually up to the apparent norm for
the species, probably because the usual hosts have a long nestling
period and there is ample time for optimum feeding and growth
by the maggots. Of the known species, broadened surstylus in

P. beameri = 113

the males is found only in avium, asiovora, and deceptor, and the
last named is a much different species than the others, with
narrow frons and parafacial in both sexes.

P. avium (northern and northeastern) and P. asiovora
(western) are very close and adults could easily be confused.
Males are consistently different in the width of the frons and
they differ slightly in proportions of the surstylus. The immature
stages are much more distinct, asiovora having an interrupted
median ventral spine _ band. Females are virtually
indistinguishable, except by association with males and
distribution in East or West. Females of avium and sialia might
also be confused, but the parafacial of sialia is not quite as broad
as in avium and is usually flat, without the bulging appearance of
avium.

The male from Yukon Territory seems an extraordinary
extension of range for avium, but the frons: head ratio is greater
(0.17), well beyond the observed ratio for asiovora and indeed
near the upper limit of the range in avium. The possible
confusion of these two species makes it imperative that puparia
be associated with the adults and be available to assist with the
identification. Specimens of avium are known from the
Northwest Territories and Saskatchewan, so that it may be that
avium is northern and transcontinental compared with asiovora.

The puparium of avium can be confused with those of
chrysorrhoea, hirundo and sialia. P. avium differs from
chrysorrhoea in having much shorter cuticular spines; from
hirundo in having more circumstigmatal folds, more cuticular
ridges and shorter spines on the dorsal cuticule; and from sialia
by having shorter cuticular spines and much shorter prothoracic
fringe. Adults of all species are readily separated.

Protocalliphora (P.) beameri, new species
(Fig. 31, Map 4)

Protocalliphora n. sp. B.--Horning and Barr, 1970: 73 [Idaho].

Diagnosis: Sexes concolorous, monochromatic, shining
metallic blue to bluish purple; calypteres white; frons of male of
moderate width; parafacial broad in both sexes; surstylus of male
strongly narrowed.

Male. Head with parafrontal and parafacial evenly bright
gray, almost silvery microtomentose; hairs on upper half of
occiput almost entirely whitish yellow behind row of postocular

114 ~~ Protocalliphora

setae, with a few black hairs adjacent to vertical bristles,
sometimes a vague and somewhat incomplete row of black hairs
behind the postocular setae. Body shining, metallic blue to dark
bluish purple or bluish green, scutum thinly microtomentose,
demarcating three full-length shining stripes (as viewed from
behind), the microtomentum more evident anteriorly. Abdomen,
viewed from behind and especially at a low angle, whitish
microtomentose on 2nd to 4th tergites, the 5th shining.
Calypteres opaque whitish, outer rims faintly yellowish tinged.

Frons moderately broad, 0.12 times head width (0.11-0.14/10,
holotype 0.115) and 1.60 times (1.43-2.00/10) ocellar span; in
profile face prominent, fronto-facial and vibrissal angles
projecting, and broad parafacial evident; preocellar area polished,
short, equilateral acute to short acuminate; each parafrontal
slightly less than half width of the frontal vitta, distinctly
microtomentose to approximately opposite median ocellus, the
prevertical area chiefly shining black, parafrontal with one to
two rows of hairs, one row beginning opposite ocellar bristles,
becoming two rows midway and merging into the numerous
parafacial hairs, latter in about four to five irregular rows;
parafacial of moderate width, slightly wider than moderately
broad frons (1.06x; 0.92-1.28)/10), obviously much wider than
ocellar span (1.70x; 1.50-1.93/10), and 4/5 vibrissal interval
(0.82x; 0.75-0.92/10); cheek broad, over 2/5 maximum length of
eye (0.44x; 0.41-0.47/10); occiput predominantly whitish haired
behind and below postocular setae, at most only a partial row of
a few slender black hairs.

Thorax without accessory notopleural bristles and hairs on
tympanic membrane; postalar wall with a few short, fine hairs.
Fore tibia with one posterior bristle.

Terminalia: cerci narrowly acuminate, deeply notched in
posterior aspect (cf. Fig. 5b); surstylus strongly narrowed on
distal half and shorter than cerci (Fig. 31); aedeagus as usual in
the genus (cf. Fig. 6a).

Female. Concolorous with male, but mesonotum duller and
much more heavily microtomentose between the narrow, shining
stripes; abdomen slightly more heavily microtomentose than in
male. Parafrontal and parafacial almost silvery microtomentose,
with slightly changeable spot on upper part of parafacial;
prevertical area dull and evenly, heavily microtomentose; hairs on
back of head darker than in male, with two irregular rows of
black hairs behind postocular setae. Frons at vertex narrower
than an eye (0.82 times) in the one available example, 0.29 times
width of head, and four times ocellar span, widening anteriorly;

P. beameri = 115

frontal vitta with a few weak hairs mesad of frontal bristles;
preocellar area finely microtomentose; 2 proclinate and one
reclinate orbital bristles; parafacial broad, its width slightly over
half that of frons (0.54x), 2.14 times ocellar span, and 1.25 times
vibrissal interval; cheek nearly half (0.47x) maximum eye length.

Apparent normal length of adults, 9.0-9.5 mm.

Immature stages. Unknown.

Type series. Holotype male, and allotype female, Cave Creek
Canyon, Chiricahua Mts., Ariz., July 4-8, 1940 (D. G. Hall).
Types in the National Museum of Natural History [USNM].

Paratypes: Arizona: male, Flagstaff, July 8, 1941 (R. H.
Beamer) [Kans. U.]; male, Prescott, June 9, 1937 (W. Benedict)
[USNM]; male, 6 mi N of Chino Valley, Yavapai Co., July 5,
1951 (R.S. Beal, reared from swallow nest) [U. Calif., Berkeley];
male, Ramsey Canyon, 5200 ft, 15 mi S Sierra Vista, Huachuca
Mts., Apr. 10, 1967 (R. F. Sternitzky) [CNC]. California: male,
Westgaard Pass, Inyo Co., June 18, 1969 (S. R. Leffler) [Wash.
State U.]; 2 males, Pine Canyon, Inyo Co., Apr. 2, 1953 (J. W.
MacSwain) [CAS]. Idaho: 4 males, Craters of the Moon National
Monument, July 16 (two), 23, and Aug. 23, 1965 (D. S. Horning,
Jr.), the July 16 specimens collected on a window, the others in
a Malaise trap [CAS, USNM]. Oregon: male, Dixie, July 8, 1931
(R. H. Beamer) [Kans. U.]. Utah: male, Box Elder Co., July 9,
1969 (K. J. Capelle, Malaise trap) [Whitworth Colln.].
Washington: male, Rattlesnake Grade, probably Benton Co., July
18, 1950 (R. B. Spurrier) [Wash. State U.]; male, Fields’ Spring
State Park, Asotin Co., July 31, 1971 (W. J. Turner, dry ice
Malaise trap) [Wash. State U.]. Mexico: Baja California Norte:
male, Arroyo Santo Domingo, 5.7 mi E Hamilton Ranch, Apr. 23,
1963 (H. B. Leech and P. H. Arnaud, Jr.), and male, Sierra San
Pedro Martir, 3000 ft, 2 mi W Socorro, June 4, 1958 (J. Powell)
[CAS].

Non-paratype Material. One male, Craters of the Moon
National Monument, Idaho, Aug. 23, 1965 (Horning) [U. Idaho]
lacks the abdomen and has not been included in the type series.
It is labeled "Chrysothamnus nauseosus," probably meaning
collected on that plant. Another male, Santa Catalina Mts., Ariz.,
July 16, 1950 (L. D. Beamer) [Kans. U.] has the typical male
genitalia and predominantly white-haired occiput and is
therefore identified with assurance as beameri, but it is
undersized (6.5 mm) and the proportions are somewhat different
than those of more normal specimens, as is often the case with
small specimens. Most notably, the parafacial is narrower than
the frons (0.89 times) and less than 2/3 the vibrissal interval

116 = Protocalliphora

(0.64). The cheek is also much narrower (0.37) than in the
typical form. The other proportions fall within the observed
range in typical individuals.

Specimens Examined. 20 males, one female.

Distribution. Arizona, California, Idaho, Oregon, Utah,
Washington, and Mexico (Baja California).

Hosts. Only one known, a "swallow" from a nest located
beneath a road bridge over a small stream, possibly a barn
swallow.

Ecology and Biology. Unknown.

Variation. In the male, occiput chiefly whitish haired, at
most one somewhat incomplete row of black hairs on the occiput
behind the postocular setae. This seems to be unique among the
Nearctic species, males of other species having two to three often
dense rows of coarse black hairs behind the postocular setae, as
in the females. This has been consistent in the males before us,
but not in the one available female, which is associated purely on
the basis of label data identical to a male beameri. Either this is
the wrong female, or there are more black occipital hairs in
females, or this is a dark-haired variant. From its general
appearance and from the features that can be compared between
males and females, such as the breadth of the parafacial, the
female does appear to be that of beameri. Unfortunately, no
whitish occipital hairs can be used for beameri in the key for
females.

Remarks. Strongly narrowed surstylus is characteristic of
five species in the group with curved surstylus: bicolor, beameri,
brunneisquama, interrupta, spenceri (Figs. 28-32). The first
species has the 5th abdominal tergite cupreous green in both
sexes, and brunneisquama has dark brown calypteres in both
sexes, while the other species in both sexes have the abdomen
entirely bluish to bluish purple and the calypteres white. The
proportions of beameri, notably of frons and parafacial, are
different than in the other four species. In male beameri, the
wide frons and broad parafacial are actually suggestive of species
such as avium, asiovora, and chrysorrhoea, which have very broad
or straight and digitate surstylus. Females, if we have associated
the right one, have no particularly distinctive features.

The superficial similarity of the bluish males to those of other
species is evidenced by the fact that one of the males stood ina
collection identified as sialia and another as hirundo, neither
having had the genitalia dissected to reveal their characteristic
form that is quite unlike that of either sialia or hirundo.

P. bicolor 117

Etymology. The specific name, in the genitive case, is
dedicated to the memory of the late R. H. Beamer of the
University of Kansas, leader of many field trips into the
Southwest, and collector extraordinary. The holotype and
allotype were collected during a field trip by David G. Hall with
Beamer and his students.

Protocalliphora (P.) bicolor, new species
(Figs. 29, 40, 45, Map 11)

Protocalliphora splendidaform splendida(Macquart).--Johnson,
1925a: 216, in part.

P. splendida_ typical.--Johnson, 1927c: 79 [N.H., from
black-throated blue warbler]; Johnson, 1929a: 29 [N.H., from
chestnut-sided warbler]; Johnson, 1930: 171-2 [N.H., from
American redstart and solitary vireo].

Apaulina metallica, in part.--George and Mitchell, 1948:
549-550.

Diagnosis. Sexes concolorous, dichromatic, both with 5th
abdominal tergite brightly cupreous, contrasting with dark blue
to bluish green body; surstylus of male decidedly narrowed.

Male. Head with parafrontal and parafacial bright gray to
silvery microtomentose, the microtomentum thinning posteriorly
as parafrontals narrow, the prevertical area chiefly polished
black; parafacial without changeable spot; occiput with 2 to 3
irregular rows of long black hairs behind row of postocular setae.
Thorax shining, dark blue to bluish green or bluish black, thinly
microtomentose, microtomentum of scutum outlining three broad
shining stripes, as viewed from behind. Abdomen bluish to
bluish green (often the latter but specimens possibly not fully
colored), tergites 2 to 4 with thin gray microtomentum that is
most evident on 2 and 3, 4 often quite shining with the "bloom"
visible only from behind when viewed at a low angle, 5th tergite
usually bright cupreous, distinctly contrasting with the preceding
tergites, sometimes green or greenish blue tinted with cupreous.
Calypteres opaque whitish, outer rims slightly yellowish.

Frons (Fig. 40) narrow and narrowing slightly anteriorly, at
narrowest only 0.065 times (0.05-0.08) head width and slightly
broader than ocellar span (1.08x; 0.83-1.40); parafrontal strongly
narrowed posteriorly, frontal vitta over twice width of a
parafrontal, polished preocellar area narrow and acuminate; lower
third to half of parafrontal with one row of hairs, or irregularly

118 Protocalliphora

two rows, parafacial sparsely haired, with 18-20 hairs in two or
three irregular rows; parafacial wider than breadth of 3rd
antennal segment, much wider than frons (1.67x; 1.29-2.00), 1.81
times (1.50-2.10) ocellar span, and 0.71 times (0.615-0.81)
vibrissal interval; breadth of cheek nearly 2/5 greatest length of
eye (0.38x; 0.36-0.40). |

Thorax without accessory notopleural bristles; tympanic
membrane and postalar wall usually bare. Fore tibia with a single
posterior bristle.

Terminalia: cerci narrowly acuminate (as in Fig. 5b); surstylus
elongate and strongly narrowed, in profile its sides subparallel on
most of its length (Fig. 29); aedeagus as usual for the genus (as in
Fig. 6a).

Female. Parafrontal and parafacial with microtomentum
grayish to weakly brownish yellow, not silvery as in male, and
likewise without changeable spots; thorax and abdomen, except
5th tergite, dark bluish to bluish green, quite dull compared with
male, heavily gray microtomentose, Sth tergite shining cupreous;
scutum, viewed from behind, with three shining stripes well
separated by heavily microtomentose areas.

Frons narrowed above, width at vertex narrower than usual
in genus, obviously much less than width of eye in dorsal aspect,
about 1/4 head width (0.24x; 0.22-0.26), and 3.29 times ocellar
span (3.00-3.83); frontal vitta with only fine hairs toward the
sides, and twice width of a parafrontal, preocellar area
microtomentose, sometimes thinly so and subshining; parafrontal
narrowing slightly posteriorly, microtomentose throughout but
thinly so and darkly subshining in prevertical area; parafacial
moderately setose, in two to three irregular rows, its width half
as wide as frons (0.50x; 0.425-0.57), 1.65 times (1.38-2.00) ocellar
span, and 0.77 times (0.65-0.86) vibrissal interval; breadth of
cheek 0.38 times (0.36-0.42) maximum length of eye.

Apparent normal length of adults, 8.5-9 mm.

Third instar. Peritreme closed; spines numerous between
stigmatal plates and in hypostigmatal area. Prothoracic fringe 82
lt (31-165)/110. Cephalopharyngeal skeleton 845 w (715-960)/41
in length.

Puparium. Length 6.9 mm (4.5-9.0)/25, breadth 3.2 mm
(2.3-3.6)/25; thin walled, dull, light brown. Posterior region (Fig.
45a, b): stigmatal plates 170 uw (140-200)/76 in diameter; distance
between buttons 505 u (330-660)/38, and across stigmatal plates
1080 ww (730-1250)/38; stigmatal ratio 0.47 (0.42-0.55)/38;
stigmatal area outlined in plaques; mesostigmatal spines scarce;
mesostigmatal folds and plaques present, markedly separating the

P. bicolor 119

stigmatal plates; hyperstigmatal spines slender, 30 wu (14-58)/215;
hypostigmatal area with numerous, weak, irregular folds;
hypostigmatal spines numerous, 6 wf (4-31)/215; circumstigmatal
folds weak and concentric. Dorsal cuticle (Fig. 45d) with
numerous spines, 25 w (12-41)/215; cuticular ridges weak or
absent. Ventral cuticle (Fig. 45c): ventral band ratio 0.55
(0.50-0.65)/50; spines of anterior patch 15 pw (6-31)/215; medial
band pronounced, spines 21 wu (10-47)/215; posterior band
pronounced.

Type Series. Holotype male, allotype, and 5 males, 20
females paratypes, Ashland, N.H., reared Aug. 4-9, 1927 by C.
W. Johnson, from nest of black-throated blue warbler collected
by Mrs. R. B. Harding. Type and allotype at MCZ through the
courtesy of Prof. A. G. Humes of Boston University, paratypes
at Boston U., USNM, and Field Mus. (1 male). Additional
paratypes: Canada: Ontario: male, two females, Algonquin Park,
July 1954 (G. F. Bennett), from yellow warbler nest; male, same
locality, October 1954 (Bennett), from nest of eastern kingbird;
3 males, 5 females, same locality, 1955 (Bennett), from nest of
myrtle warbler [all USNM]. Contiguous United States: New
Hampshire: 2 males, Ashland (Mrs. R. B. Harding), from nest of
chestnut-sided warbler [Boston U.]; male, female, Ashland, July
20, 1929 (Mrs. Harding, from nest of black-throated blue warbler
[CNC]; male, Ashland, August 1927 (Mrs. Harding), nest of
black-throated blue warbler [USNM]; 6 males, 4 females,
Holderness, July 18, 1919, from black-throated blue warbler
[MCZ]; 3 males, 3 females, Holderness, July 20, host not stated
[MCZ]; 5 males, 5 females, Holderness, Aug. 10, 1930, nest of
American redstart (Mrs. Harding) [MCZ]; 10 males, 6 females,
same locality, host, and collector as preceding, but nest collected
June 27, 1930, each specimen with puparium [Boston U.]; 2
males, 2 females, Holderness, Aug. 10(?), 1930 (Mrs. Harding),
from nest of solitary vireo [MCZ]; 19 males, 20 females,
Holderness, July 15, 1929 (Mrs. Harding), from nest of solitary
vireo [MCZ]; male, Rumney, Dec. 22, 1924 (P. J. Darlington),
"chopping dead pine" [Boston U.]; 2 males, Grafton Co.,
Livermore, East Pond, Aug. 23-24, 1980, on Solidago (J. F.
Burger) [U. of New Hampshire]. New York: male, Lake Clear
Junction, Franklin Co., Aug. 10, 1946 (J. L. George, R. T.
Mitchell), with puparium [USNM], mixed with Apaulina
metallica of George and Mitchell (1948); male, Whiteface Mt.,
Essex Co., Sept. 13, 1979 (A. Freidberg) [USNM]. Wisconsin: 9
males, 5 females, Madison, with puparia, from nest of alder
flycatcher, July 1974 [U. Wis., USNM].

120 ~=Protocalliphora

Non- paratype Material: A male and two females from various
New Hampshire lots, and 2 males, 2 females, from Algonquin
Park, Ont., from nest of eastern kingbird, are teneral and have
not been included as paratypes. Puparia identified as bicolor
were found by Bennett at Algonquin Park in nests of common
grackle, American tree swallow, wood thrush, and red-eyed
vireo, and at Toronto, Ont., in nest of American goldfinch, but
no adults were reared. Two isolated females, from Peterboro,
N.H., Aug. 29, 1924 (C. F. Batchelder) and Little Black River
Rapids, Maine (near mouth of Allegash River, upper Aroostook
Co.), Sept. 8, 1907 (J. A. Cushman) [Boston U.] are probably
bicolor, having narrow frons, broad parafacials, and long third
antennal segment. At least the second of these was recorded by
Johnson (1925: 216) as Protocalliphora splendida form splendida.

Two females, Val d’Or, Que. [CNC], each mounted with
puparium, definitely are bicolor, but there may have been some
confusion in labeling. The abbreviated labels signify "Ex
Estigmene acraea (Arctiidae), emerged 7-29-1942, Recherches
des insectes forestiers," but the association with the lepidopteran
is certainly incorrect.

Specimens Examined. 77 males, 83 females.

Distribution. Northeastern: Wisconsin, Ontario, New York,
and New Hampshire, and probably Quebec and Maine.

Basis of description of immature stages: Larva, 3rd instar (7
specimens): Algonquin Park, Ont., yellow warbler, July, 1954 (6),
and myrtle warbler, July, 1955 (1). Puparium (38 specimens,
partly from type series): Algonquin Park, Ont., yellow warbler,
July, 1954 (22) and American goldfinch, August, 1954 (2);
Ashland, N.H., black-throated blue warbler (9); Holderness, N.
H., solitary vireo (2), and American redstart (3).

Hosts. Alder flycatcher, eastern kingbird, American redstart,
solitary vireo, and various warblers (black-throated blue,
chestnut-sided, myrtle, and yellow); also, recorded from puparia
alone, American goldfinch, common grackle, American tree
swallow, wood thrush, and red-eyed vireo.

Ecology and Biology. This species has been taken in only a
few localities, chiefly from birds nesting in the intermediate
levels of deciduous forests. If bicolor occurs characteristically in
this habitat it may be more common and more widely distributed
than the records indicate because the habitat is less thoroughly
explored than others and is more difficult to survey.

Remarks. Protocalliphora bicolor is unique among the eastern
species in having the surstylus so strongly narrowed. Several
western species have equally narrow surstylus, but they are easily

P. bicolor 121

distinguished from bicolor by the entirely blue abdomen of the
males. Females of bicolor fall in the group of species with
abdomen predominantly bluish but fifth tergite cupreous, the
other species being metallica and the western cuprina. These will
be difficult to separate satisfactorily, except when males and/or
immature stages are associated with them, and isolated females
have usually not been positively identified in this study. In the
East, where females of bicolor and metallica can easily be
confused, the puparia are very distinctive. The integument in
bicolor is well covered with distinct spines, whereas in metallica
the cuticular spines are fine and minute, and the dorsal plate
appears to be finely granular, with a sandpaperlike appearance.
Furthermore in females of bicolor, the parafacial has no
changeable spot, whereas the other two species have an
arrangement of microtomentum on the upper part of each
parafacial that results in a large dark spot on a light parafacial,
viewed from above, changing to a light spot on a dark field when
viewed from below.

Puparia of bicolor are easily confused with those of cuprina
and parorum. They are separated with difficulty from cuprina on
the basis of the somewhat longer prothoracic fringe and short
tubercle-like spines of the hypostigmatal area and from parorum
on the basis of the smaller ventral band ratio and shorter
tubercle-like spines of the hypostigmatal area. This species has
thus far only been recorded in the eastern portion of the
continent while the other two are western in distribution.

Teneral males of bicolor often have the fifth tergite only
slightly cupreous, and these may be confused with teneral males
of metallica in which the fifth tergite is occasionally somewhat
greenish tinted. Male terminalia or puparia will easily settle this
question of identity.

The holotype and specimens reared with it were recorded as
typical P. splendida by Johnson (1927c: 79), who reared a total
of 9 males, 26 females from the nest on Aug. 4, 5, and 9, 1927.
Johnson remarked that the abdomen of both sexes showed a
coppery fifth tergite but he believed that this merely refuted
Shannon and Dobroscky’s (1924) belief in sexual dichromatism
for splendida. However, their specimens were correctly
interpreted (cf. discussion under metallica), and Johnson’s
specimens represented an undescribed species, here described as
bicolor. Both series from the American redstart and that from
the solitary vireo (called in Johnson the blue-headed vireo) were
recorded as typical splendida by Johnson (1930), and the Ashland

122 Protocalliphora

specimens from the chestnut-sided warbler were recorded as the
same by Johnson (1929a: 29).

The small series from the myrtle warbler and the 1929 series
from the solitary vireo contain many small and certainly
undersized specimens, ranging from 5-8 mm in length. In these
small specimens, the fifth tergite is usually only slightly tinted
with cupreous, not strongly cupreous as in more typical
specimens. Some of these are teneral, which would also affect
the development of mature color.

Etymology: The specific name is a Latin adjective, referring
to the two colors of the abdomen, and invariable.

Protocalliphora (Trypocalliphora) braueri (Hendel)
(Figs. 6c, 20, 41, 46, Map 3)

Avihospita braueri Hendel, 1901: 30 [Provenance not stated, later
known to be Austria].

Protocalliphora chrysorrhaea [sic] (Meigen).--Walton, 1914: 175,
in part [N. Mex., later paratypes of hirudo].

P. azurea (Fallén).--Arnold, 1919: 147-8 [Colo.; paratypes of
hirudo].

Phormia metallica Townsend.--Plath, 1919c: 376 [Wash.; became
type series of Protocalliphora hirudo var. cuprea]; Townsend,
1919: 379-80 [Wash., atypical form of metallica, also proved
to be hirudo cuprea}.

Protocalliphora azurea.--Cole and Lovett, 1921: 309 [Oreg.].

P. hirudo, P. hirudo var. parva (subspecies?), and P. hirudo var.
cuprea Shannon and Dobroscky, 1924: 249, 252-3 [hirudo,
Colo., N. Mex.; parva, Kans.; cuprea, Wash.].

P. hirudo var. cuprea.--Johnson, 1925a: 216 [Mass.]; Johnson,
1925b: 51-3 [Mass.].

P. hirudo and var. parva.--Shannon, 1926: 128 [In key].

P. splendida.--Bergtold, 1927: 106-7 [Colo.; larvae in abscess,
undoubtedly braueri].

P. hirudo cuprea and P. sp. on baby chicks.--Hearle, 1938: 60-1
[Undoubtedly braueri].

P. sp. larvae in golden eagle nest.--Philip, 1938: 486-7 [Alaska].

P., hesperia.--Guberlet and Hotson, 1940: 65-8 [Wash.].

Apaulina hirudo.--Hall, 1948: 192-5 [Revision; varr. cuprea and
parva possibly synonyms; Kansas as type locality for parva
corrected to Greeley, Colo.]; George and Mitchell, 1948:
550-2 [N.Y.].

Protocalliphora hirudo.--Munro, 1949: 35-38 [B.C.].

P. braveri = 123

P. sp. or spp.--Wood, 1953: 77 [B.C.; probably braueri in part);
Dodge and Seago, 1954: 53-4 [Ga.].

Trypocalliphora hirudo.--Peus, 1960: 225 [Europe; reference to
new genus, hirudo and braueri considered distinct species].

Protocalliphora hirudo.--Hall, 1965: 926 [Catalog; widespread;
parva and cuprea synonyms].

P. hirudo.--Cais, 1965: 183-191 [Poland; good figures, male
genitalia and larvae].

P. (Trypocalliphora) hirudo.--Sabrosky, 1967: 122
[Trypocalliphora accepted as subgenus rather than genus].

P. hirudo.--James, 1969: 252 [Mention of biology]; Rausch, 1972:
1-4 [Alaska]; Whitworth, 1976: 16 ff.; Whitworth, 1977:
4933-B [Mention]; Bedard and McNeil, 1979: 111 [Quebec].

Trypocalliphora braueri.--Rognes, 1985: 373 [Europe; detailed
description and notes on type series from Austria; hirudo a
synonym].

Diagnosis.--Conspicuously dichromatic, the thorax and
abdomen shining, metallic bluish black in male, aeneous green
and only weakly shining in female; accessory notopleural bristle
almost always present, rarely absent on both sides; male with
extremely narrow frons and uniquely characteristic surstylus and
aedeagus (Figs. 6c, 20); female lacking reclinate orbital bristles,
and with narrow frons.

Male.--Head with microtomentum of parafrontal and
parafacial yellowish to brownish tinted. Thorax shining dark
bluish black, or with slight greenish-blue tint, the
microtomentum thin and fine, usually evident only on anterior
slope of scutum; fringe of mesothoracic spiracle orange-yellow
to brownish, not as bright as in Phormia regina but brighter than
in any other Protocalliphora. Abdomen dark blue to bluish
black, highly shining, with thin gray "bloom" evident only at a
low angle from behind. Calypteres yellow to brownish tinted,
perhaps typically light brown in mature specimens.

Frons narrow (Fig. 41), obviously narrowing anterior to
ocelli, at narrowest 0.05 times head width (0.04-0.07, with 19 of
the sample between 0.04 and 0.06), and narrower than ocellar
span (0.73x; 0.60-0.93); parafrontals clearly separated, though
narrowly, the frontal vitta visible throughout, subequal to or
wider than a parafrontal and widening strongly anterior to the
narrowest point; preocellar area narrow, acuminate, rugose,
shining and not microtomentose; parafrontal narrow and
microtomentose below, with a single row of hairs outside the
frontal bristles, above strongly narrowed and_ thinly

124 Protocalliphora

microtomentose or bare, shining black; parafacial narrow, densely
microtomentose, with two to three irregular rows of black hairs,
weakly ridged, width of each parafacial only a little wider than
ocellar span (1.31x; 1.07-1.60), wider than narrow frons (1.79x;
1.31-2.28), and not 2/3 vibrissal interval (0.64x; 0.56-0.75/23);
frontal bristles rapidly decreasing in size above, the uppermost
short and hairlike, rows ending well before median ocellus, and
approximately opposite apex of the acuminate preocellar area;
cheek comparatively narrow, its maximum height 1/3 maximum
length of eye (0.35x; 0.32-0.39).

Thorax: Accessory notopleural bristle almost invariably
present, rarely absent and then usually on one side only (cf. Fig.
3a); tympanic membrane either bare or haired, often with strong
black hairs (For both sexes, entirely bare in 43 specimens, but
hairs on one or both sides in 28, ranging from a single pale weak
hair to 13 coarse black hairs); postalar wall bare, rarely with a
weak adventitious hair. Fore tibia with one median posterior
bristle.

Terminalia distinctive: cerci straight, long and slender (Fig.
20); surstylus with characteristic profile (Fig. 20), almost
boot-shaped, broad at base, narrowing strongly, then slightly
broadened distally and tending to curve anteriorly; distal portion
of aedeagus cylindrical, distal margin flared (Fig. 6c).

Female. Body aeneous green, rather heavily microtomentose
and only weakly shining, scutum usually with the three shining
stripes narrow but more or less evident, abdomen more shining
than thorax but under microscope obviously densely and finely
microtomentose; tergite 5 often especially shining and cupreous
green. Calypteres whitish yellow to orange yellow (typical), paler
than in male, rims dark yellow.

Frons extremely narrow for female Protocalliphora, half or
less width of an eye, less than 1/4 width of head (0.22x;
0.21-0.25), and less than three times ocellar span (2.86x;
2.43-3.38); parafrontal narrow, dull, gray to brownish gray
microtomentose, prevertical area dull and_ densely
microtomentose; frontal vitta with few weak and inconspicuous
hairs, preocellar area usually heavily brown-gray
microtomentose, contrasting with brown of frontal vitta,
occasionally thinly microtomentose and more shining; reclinate
orbital bristles absent; parafacial narrow, its width less than half
width of frons (0.44x; 0.36-0.50), only 1 1/4 times ocellar span
(1.25x; 1.07-1.43), and 2/3 vibrissal interval (0.67x;
0.57-0.82/22); cheek comparatively narrow, 1/3 maximum eye
length (0.34x; 0.30-0.37).

P. brauert = 125

Thorax and legs as in male.

Apparent normal length of adults, 7.5-8.5 mm.

Second instar. Cephalopharyngeal skeleton 370 wu in length.
Prothoracic fringe 8 ub (4-15)/15; spines scarce. Posterior region:
Peritreme open, button indistinct; stigmatal plates 71 mw in
diameter; mesostigmatal spines absent; hyperstigmatal spines
scarce, 7 wt (2-14)/30; hypostigmatal spines absent. Dorsal
cuticle with narrow band of spines, one-fourth width of segment,
on anterior margin of each segment, passing ventrally as the
anterior band, spines 9 uw (4-18)/30. Ventral cuticle with single
(anterior) band of spines, 9 u (4-18)/30; spines of anterior patch
scarce and small, 5 uw (2-8)/30; medial and posterior bands
absent.

Third instar. Peritreme usually open; spines absent from
meso- and hypostigmatal areas. Prothoracic fringe 7 yt (3-10)/60.
Cephalopharyngeal skeleton 734 w (560-860)/23 in length.

Puparium. Length 7.2 mm. (4.3-8.3)/26, breadth 3.2 mm.
(1.9-3.9)/32; thin walled, semi-translucent, shiny, light brown.
Posterior region (Fig. 46a, b, e): Stigmatal plates 188 wu
(130-230)/90 in diameter, distance between buttons 422 wu
(260-545)/46, and across stigmatal plates 1050 wu (675-1420)/46;
stigmatal ratio 0.40 (0.32-0.46)/46; stigmatal area often outlined
in plaques; mesostigmatal spines absent; mesostigmatal folds
and/or plaques present; hyperstigmatal spines extremely scarce,
6 pw (3-12)/120, sometimes absent; hypostigmatal area with
pronounced, irregular folds (western material), or folds weak or
absent (eastern material); circumstigmatal folds variable, in
western material prominent and irregular or rectangular (Fig.
46a), in eastern material weak and rectangular, or absent (Fig.
46e). Dorsal cuticle with spines or scarce, 8 ww (4-12)/130;
cuticular ridges variable, pronounced, weak, or absent. Ventral
cuticle (Fig. 46 d, f): ventral band ratio 0.11 (0.08-0.25); spines
of anterior patch not distinguishable from other spines of
anterior band, 9 uw (4-17)/200; medial band absent; posterior
band sometimes seen, normally absent; each segment usually
subdivided by at least one annulus between the scars of the
ventral pads.

Type Series. P. braueri: The type series, from Austria
[Naturhistorisches Museum, Wien], has been reported on in detail
by Rognes (1985).

P. hirudo: Holotype male and allotype, Colorado, July 10,
1911 (M. A. Palmer), "ex nesting warbler"; paratypes, male, 2
females, Koehler, N. Mex. (W. R. Walton, Webster No. 7707), and

126 _—_ Protocalliphora

male, 3 females, Colorado Springs, Colo., August 1916 (W. W.
Arnold), "from maggot in sparrow a[nd] goldfinch" [last

3 females in MCZ]. One of the female paratypes from Koehler,
N. Mex., has not been located. [All USNM except as noted.].

P. hirudo var. or subsp. parva: Holotype male and allotype,
"Kansas," but correctly Greeley, Colo. [USNM].

P. hirudo var. cuprea: Holotype and 2 paratypes, all females,
published as Seattle, Wash., labeled as Puget Sound (O. E. Plath),
from nest of "western" [= American] robin. [USNM].

Under typical hirudo in the USNM collection are two other
females with data like the Koehler, N. Mex., paratypes. One of
these is also labeled "ex horned lark Otocoris alpestris." These
Koehler specimens are part of the 30 specimens of
"Protocalliphora chrysorrhaea" [sic] reported by Walton (1914:
175), which was a mixed infestation (See P. seminuda n. sp. for
a part of this series).

The type material of P. hirudo parva was twice recorded in
error, according to information in J. M. Aldrich’s card catalogue
of Diptera [USNM]. Aldrich, on a card then headed
Protocalliphora chrysorrhoea, noted that "Williston gave me two
pairs that he got from a man in Kansas, who found the larvae
under the skin on the back of the head in young birds; I labeled
them ’Young birds, Kans., S.W.W." Shannon and Dobroscky,
having one male and one female of this lot (the former headless
and the latter without abdomen!) described P. hirudo parva as
"reared from fledglings, Kansas, (S.W. Williston through Prof. L.
L. Adams)." Aldrich thereupon noted on his card for parva: "This
should be reversed, it really was Greeley, Colo. (L. L. Adams,
through S. W. Williston)," and he placed a label in the collection,
"Prof. L. L. Adams, Greeley, Colo. Ex nestlings." Hall (1948: 195)
apparently missed the Aldrich notes, and credited collection of
the specimens to Williston, although correctly recording the type
locality as Greeley, Colo. Incidentally, the specimens of parva
are undersized individuals, and the male holotype lacks distinct
accessory notopleural bristles, as noted by Hall. The allotype has
an accessory notopleural on the left side only.

The variety cuprea was apparently based on three females,
the holotype and a paratype deposited in the U.S. National
Museum and a female paratype that came later with the Shannon
Collection. A fourth female from the same rearing is teneral,
with ptilinum extruded, and was not labeled as a paratype. These
are the "four Puget Sound specimens" included by Townsend
(1919) in the type series of his Phormia metallica.

P. braveri = 127

Specimens Examined. 121 males, 137 females. If a nest is
not specified, larvae were taken directly from the bird noted.
102 males, 114 females were from the West, the balance from the
East.

Distribution. Canadian and Transition Zones: in the West,
Alaska to Saskatchewan, south to Mexico (Durango), New
Mexico, and Colorado; in the East, Ontario and Michigan to
Labrador and Virginia, also Georgia.

Alaska: Mentasta Lake, Mentasta Road (Wilson’s warbler);
Rapids, S of Big Delta (golden eagle nest); Steese Highway, mile
105 (American tree sparrow nest).

Canada: Alberta: Edmonton (house wren). British Columbia:
Beaver Lake, SW of Likely (young bird); Bella Coola (junco nest);
Kamloops (nests of Brewer’s blackbird, cliff swallow, and
western kingbird); Lilloet, Mt. McLean (chipping sparrow);
Prince George (house sparrow nest); Qualicum (barn swallow);
Terrace; 6 mi W Terrace; 32 mi S Terrace; Vancouver (barn
swallow nest, house sparrow, western meadowlark, Swainson’s
thrush); North Vancouver (budgerigar); Vanderhoof, W of Prince
George (song sparrow); Vernon; Victoria (barn swallow).
Labrador: Cartwright (white-crowned sparrow). Newfoundland:
Codroy Valley, near Port-aux-Basques (American robin nest).
Ontario: Algonquin Park (great crested and alder flycatchers,
vesper sparrow, and Canada and myrtle warblers, and in nests of
common grackle, ovenbird, vesper sparrow, barn and bank
swallows, and yellow warbler); Ogoki. Prince Edward Island:
Cavendish Beach, Prince Edward Island National Park. Quebec:
Isle Verte, 15 km E of Riviére-du-Loup (savannah sparrow); La
Verendrye Provincial Park; Val d’Espoir. Saskatchewan:
Muscow, S of Fort Qu’Appelle (wren).

Contiguous United States, West: California: Columbia;
Fresno Co. (Bear Creek); 8 mi E Georgetown (mountain
chickadee); Tuolumne Co. (black-headed grosbeak). Colorado:
"Colo." (warbler); Colorado Springs (sparrow and goldfinch); near
Fort Collins (loggerhead shrike); Fort Collins; Greeley. Idaho:
Lake Waha; Lewiston. Montana: Flathead Lake, Lake Co.
(Brewer’s blackbird); Hamilton (sparrow); Ravalli Co.
(violet-green swallow, bluebird nest); Sunburst, Toole Co.
(sparrow, probably vesper). Nevada: Washoe Co. (house wren).
New Mexico: Abique Dam, Rio Arriba Co. (cliff swallow nest);
Koehler (horned lark). Oregon: Lents, now in Portland (young
bird); Milwaukie (tree swallow). Utah: Mantua in Box Elder Co.
(house wren, subcutaneous). Washington: Olympia (barn
swallow); Puyallup (house sparrow); Seattle (yellow warbler, and

128 Protocalliphora

in nests of American robin and a sparrow); Tacoma (barn
swallow). Wyoming: Laramie (house sparrow); Moran (pine
siskin).

Contiguous United States, East: Georgia: Rabun Co., Rabun
Bald at summit, 4714 ft. Massachusetts: near Middleboro
(sparrow). Michigan: Iron Co., Gaastra ("ground sparrow"). New
York: Franklin Co., Lake Clear Junction (chipping sparrow).
Virginia: Giles Co. (great crested flycatcher); Roanoke (house
wren); larvae only, Augusta Co. (loggerhead shrike).

Mexico: Male, Durango, 11 mi W Durango, June 20, 1964 (J.
F. McAlpine) [CNC]; female, Rio Chico, 20 mi W Durango, July
22, 1964 (J. F. McAlpine) [CNC].

Most available specimens were reared June-August. An early
record was the rearing of three males, three females at Puyallup,
Wash., Apr. 28, 1926, "from larvae on Passer domesticus" (Theo.
H. Schaffer).

Basis of description of immature stages: Larvae, 2nd instar
(1 specimen): Algonquin Park, Ont., ovenbird, June 1954. Third
instar (9 specimens, plus some measures of prothoracic fringe and
cephalopharyngeal skeleton from western puparia): Algonquin
Park, Ont., ovenbird, June 1954 (4), and yellow warbler, June
1954 (5). Puparium, western (25 specimens): Hamilton, Mont.,
"sparrow" fledgling, July 1936 (10); Kamloops, B. C., Arkansas
(i.e., western) kingbird, August 1945 (4); Vanderhoof, B. C., song
sparrow, July 1945 (7); Lilloet, B. C., chipping sparrow, June
1947 (2); Lulu Island, Vancouver, B. C., western meadowlark,
June 1932 (2).

Puparium, eastern (21 specimens): Algonquin Park, Ont.,
ovenbird, June 1954 (10), myrtle warbler, June 1945 (6), and
Canada warbler, June 1945 (2); "Ontario" (Halket), "sparrow" (1);
Lake Clear, N.Y. (2).

Hosts. Western: Brewer’s and yellow-headed blackbirds, a
bluebird (Montana, Oregon), mountain chickadee, golden eagle,
house finch, American goldfinch, black-headed grosbeak, a
junco (probably Oregon junco, from north of Vancouver, B.C.,
in breeding range of Oregon junco), western kingbird, horned
lark, black-billed magpie, western meadowlark, American robin,
loggerhead shrike, pine siskin, Townsend’s solitaire (probably;
Munro 1949), sparrows (chipping, house, song, American tree,
and probably vesper), European starling, swallows (barn, cliff,
tree, and violet-green), Swainson’s thrush, warblers (Wilson’s,
yellow, and unidentified warbler in Colorado), and house and
long-billed marsh wrens, also budgerigar and possibly baby
chicks (Hearle 1938).

P. braveri 129

Eastern: Great-crested and alder flycatchers, common
grackle, ovenbird, American robin, loggerhead shrike, sparrows
(chipping, savannah, vesper, white-crowned, and "ground"), bank
and barn swallows, warblers (Canada, myrtle, yellow), and house
wren.

Ecology and Biology. Presumably ubiquitous, judging from
the recorded hosts, but more frequent in nests or on birds at
ground or shrub level--or young birds are more often observed
closely in these situations! Maggots of this species are often
found subcutaneously, causing true myiasis.

The myiasis-producing habit of larvae of P. braueri was
mentioned when its synonym, hirudo, was first described,
although it was not then realized how characteristic that was
because some specimens were reared from puparia found in
nests, as is normal for other species of Protocalliphora. Shannon
and Dobroscky (1924) described both typical hirudo and hirudo
parva from maggots found in nestlings (cf. Type Series), but they
did not give details. They also recorded as "aenea" a fragmentary
specimen, now recognized as braueri, reared from a larva "from
brain of living fledgling of sparrow kind." However, Walton
(1914) and Arnold (1919) had already published similar
observations on myiasis. Walton stated that larvae of what he
called P. "chrysorrhaea" [sic] on a fledgling horned lark "were
contained in purulent sores on the sides of the body near the legs
and on the neck." (N.B. The infestation was a mixed one; see
discussion under seminuda). Arnold (1919), writing of P.
"azurea" in his article on "Maggot infested birds," stated that a
large number of young birds of various species were infested
with maggots, and that "the young [larva] immediately burrows
beneath the surface, -- is on the head, near or in the corner of
the eyes, about the neck and upper part of the wings....The
maggot eats a burrow or chimney into the flesh of its victim,
remaining stationary and feeding upon the fluids of the helpless
victim until death releases it." Two young meadowlarks attacked
by a hundred maggots had bodies "presenting the appearance of
having been struck by a load of shot."

A further observation of special interest is that of R. C.
Miller (1929). Maggots emerged from a young yellow warbler
(Dendroica aestiva, now petechia) barely able to balance on a
twig. Dissection after the bird’s death showed the following:
"Eleven subcutaneous pockets that had contained one or more
larvae were found, three on the head, three on the neck, and the
remainder scattered over the body, especially under the wings
and on the breast. Two of the lesions entered the pleural cavity,

130 — Protocalliphora

and the lungs had been partly devoured." The author concluded
that this seemed definite evidence that when larvae were "present
in sufficient numbers they may literally devour the young birds
alive." In all 23 larvae emerged or were removed from the bird.
A few reared adults were identified by J. M. Aldrich as
Protocalliphora splendida var. hesperia, but a male and female
preserved in the National Museum of Natural History in
Washington show that the species is braueri.

After the Shannon and Dobroscky revision (1924), Johnson
(1925) recorded the rearing of adults of hirudo cuprea [male,
female, now in Boston U.] from maggots in the neck of a
sparrow, and in 1931 he added that these had been found in
"swellings on the neck." Bergtold (1927) recorded P. "splendida"
as reared from a young house finch at Denver, Colo., from larvae
in an abscess in a good-sized swelling below the right eye, and
we believe that this record almost certainly refers to braueri. Still
later, Guberlet and Hotson (1940) gave details of the myiasis
observed in a young house sparrow at Seattle, Wash., from which
flies identified as "P. hesperia" were reared [A male from this
rearing has been found to be braueri]. After the death of the
young bird, 21 live maggots were removed from the skin and
subcutaneous tissue on the sides of the body, the neck, the right
wing, and above the right eye. They were in slight swellings,
which had small pores at which were located the anal ends of the
larvae. George and Mitchell (1948), working at Lake Clear
Junction, N.Y., in 1946, reared 3 males, 3 females of "hirudo"
[USNM] from nestling chipping sparrows, and recorded that 5 of
the 12 maggots observed were completely embedded in the skin
of one nestling, which died. Munro (1949) found hirudo larvae
abundant, feeding subcutaneously and causing mortality up to 75
%. Rausch (1972) described a case of subcutaneous myiasis in a
fledgling Wilson’s warbler in Alaska, with descriptions of both
macroscopic and microscopic characteristics of the lesions. The
larvae remained in the subcutaneous cavities until the death of
the bird. A recent note by Bedard and McNeil (1979) recorded
detailed observations on the larvae of braueri (as hirudo) on
savannah sparrow nestlings (four in one brood in 1976, three in
another brood in 1977): "The seven nestlings each supported
upwards of 10 fly larvae and the site[s] of embedment, in
decreasing order of importance were (a) the wing tracts
(especially between the base of the remiges), (b) the head
including the nasal cavity, the ear duct, and the area close to the
eyes, (c) the crural and abdominal tracts, and (d) the legs. One
nestling had three larvae moving freely between the naris and a

P. braveri —s 131

breathing pore on the nape. All larvae were embedded
subcutaneously and vigorously resisted extraction, suggesting that
P. hirudo is an obligate subcutaneous parasite... The presence of
these parasites retarded feather growth and the plumage around
the site of the embedment had a sticky appearance. One of the
nestlings lost an eye due to a parasite larva." The incidence of
parasitism was low; one brood of 25 examined in 1976, one of 34
in 1977.

Brief notes on the labels of specimens submitted for
identification sometimes state the subcutaneous location of larvae:
male, three females, Lilloet, B.C., July 24, 1947 (H. C. Coppel),
"internal in Spizella passerina " [UBC]; two males, three females,
Ravalli Co., Mont., July 1967 (Sheri Marino and W. L. Jellison),
"reared ex larvae in cysts of Tachycineta thalassina lepida"
[USNM]. At Algonquin Park, Ont., Bennett found a
subcutaneous larva in a young fledgling alder flycatcher, and
three larvae of braueri in an open lesion above the eye of a young
vesper sparrow. He identified as "hirudo" larvae causing myiasis
in young sparrows in Quebec City (J. McNeil); at Lents, Oregon,
two maggots came out of the head of a young bird about to fly,
and one began boring into the hand of the collector, H. A.
Darnell [two females, USNM and Oregon State U.]; four males,
two females, reared from maggots taken June 20, 1970 from a
nestling loggerhead shrike near Fort Collins, Colo. (T. J. Cade)
[Cornell U.] (These maggots were completely under the skin,,
mostly in the wings between areas of developing primary and
secondary feathers, but one under an eye, one in the back of the
neck, and some in the nares; none of the birds died).

At Roanoke, Va., Aug. 8, 1972, five young house wrens were
observed by C. H. Lewis as they emerged from the nesting box.
The feathers on the heads did not appear normal and, having
caught two of the birds, he found and removed five larvae from
under the skin of one bird and two from the other. Two
submitted as puparia for identification proved to be braueri;
presumably the others were also [Va. Polytechnic Inst.]. One
adult male with puparium, and two larvae, came from Edmonton,
Alta., July 1975 (Bruce Bembridge) [U. of Alberta]; the 11-day
old wren nestlings were said to be "afflicted with boils." At
Qualicum, B.C., R. A. Cannings recorded larvae in barn
swallows, "found under the skin of the nestlings, in the wings and
head." [BC Provincial Mus. (now Royal BC Mus.), USNM]. In
Giles Co., Va., B. A. Garrison found larvae in cysts about the
head, wings, and beak of a nestling great crested flycatcher [Va.
Polytechnic Inst., USNM].

132 Protocalliphora

Observations more detailed than usual were recorded by H.
W. Prescott at Milwaukie, Oregon (personal communication).
These apply to braueri; adults of this species were reared from
the material in the observed nest. In one of his bluebird nest
boxes, he found "a tree swallow nest with four swallow nestlings
about five days old and still essentially naked. On the top of the
head of one of the nestlings I noted a small round wound slightly
less than one sixteenth of an inch in diameter. On looking more
closely I noted that this wound was actually a hole in the
nestling’s integument filled flush to the exterior surface by a fly
maggot. At that point I inspected the other nestlings closely but
found no more lesions of any sort on them. But to my surprise
I found something else equally interesting. Each nostril of every
one of the four nestlings was plugged completely shut by a fly
larva, the truncate end of all larvae coming flush with the
exterior of the nostril surfaces. Six days later, I returned to the
swallow’s nest. The nestlings were about half grown by then and
I saw no further signs of maggots in them." So the nest was
removed--a substitute nest was provided--and the nest material
sifted for puparia from which adults of braueri emerged. He also
recalled an earlier experience that also certainly involved braueri
although no flies were reared. Among his numerous bluebird
nest boxes--a total of 470!--two "contained nestlings only a few
days old and still naked in both instances. One of the boxes
contained nestling house sparrows and the other bluebirds. In
both cases some of the nestlings had these small round wounds
above described. They seemed to occur at random on the
nestling’s anatomy. Some were on the wings, some on various
parts of the torso, and some on the legs exclusive of the tarsae,
some on the head."

When flies are reared from larvae and/or puparia found in
the nest, or when labels read simply "on...," the exact feeding
habits of the larvae cannot be determined. However, in a few
cases the details given show that the larvae of braueri do not
always live subcutaneously, or not continously. At Moran, Wyo.,
J. D. Tiner reared 8 males, 11 females "from maggots which had
anterior ends buried in craters in the skin of young pine siskin
just off nest, and spending night on ground. ... all on head, neck
and shoulders of bird." The larvae started falling off as soon as
the bird was picked up, on the night of July 16, 1951, and adult
flies emerged August 3 [USNM]. J. M. Kinsella made similar
observations ona nest of Brewer’s blackbirds near Flathead Lake,
Flathead Co., Mont.: "The birds were out of the nest in the grass,
and, when I picked them up, the larvae immediately began

P. braveri = 133

actively leaving the birds and dropping to the ground. Most of
the larvae I saw came from the neck region. The larvae pupated
in 24 hours and the adults hatched about 10 days later."

In some cases, the birds were apparently not seriously
injured, in others the young birds were said to be "weak and
emaciated" (Walton 1914). At Columbia, Calif., David C. Rentz
reared 12 males, 13 females [CAS, USNM] from "subcutaneous
larvae on a young black-headed grosbeak," and the bird died
shortly after the larvae left the body. Death was attributed to the
maggots. Damage done by maggots of this species was described
as follows by Dr. Murray L. Johnson, M.D., Tacoma, Wash.
(personal communication), who reared a specimen of "“hirudo"
[Johnson Colln.]: "I found the juvenile barn swallow alongside a
small pool of water south of Tacoma on July 19, 1947. 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
larvae were immediately put in a jar of damp earth; the swallow
died about 24 hours later." The adult fly emerged on 7 August.
Three males, Lulu Island, Vancouver, B.C., June 1932 (R. A.
Cumming) [UBC] were reared from nestling meadowlarks "very
much cut up." In the case of cutaneous myiasis by "hirudo" ina
fledgling Wilson’s warbler, described by Rausch (1972), the
larvae were in subcutaneous cavities in the head just above the
beak. The bird refused food and died the day of its capture.
Rausch concluded that "Aside from the possible adverse effects
of exsanguination, the extent of destruction of tissue was such
that the bird probably could not have survived. The lesions were
produced largely by the growth of the larvae within a restricted
locus." Under such conditions, notably infestation about the
head, Protocalliphora larvae can certainly cause great damage and
even death, or predispose to later death, or contribute to death
from a combination of factors. Cais (1965), in Poland, reached
an even stronger conclusion: "It seems definite that all observed
cases of parasitism by P. hirudo bring about the death of the
nestling, even if the number of larvae is small."

Remarks (including Variation). Protocalliphora braueri
differs from all other species of Protocalliphora in North
America in several characters that are unique in the genus: The
form of both surstylus and aedeagus in the males, the absence of
reclinate orbital bristles in the females, and accessory notopleural
bristles in both sexes. (Such bristles occur only occasionally in
other species as aberrations of chaetotaxy). The females are

134 Protocalliphora

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P. braueri = 135

further distinguished from those of all known species except
deceptor by the unusually narrow frons, and from all but females
of aenea by the aeneous to dull cupreous green body color,
contrasting with the metallic blue of most females. Unlike most
species of Protocalliphora, isolated individuals of either sex can
be readily identified, and the larvae and puparia are equally
distinct. For a discussion of the possible generic or subgeneric
significance of these characters, see the discussion on "Status of
Trypocalliphora."

In an early stage of the study, braueri (as hirudo) was
regarded as a western species, and the eastern population as a
subspecies or a narrowly distinguished sister species with slightly
narrower frons and slight differences in other proportions.
Later, after reviewing the broadly overlapping range of variation,
and also realizing the undue influence on the eastern ratios of
some undersized specimens, which had been measured because
of the relatively small numbers of available measurable examples,
we concluded that only one species was present. For possible
future use, however, we have maintained separate ratios for the
eastern and western populations. In the description, the ratios
and ranges refer to western specimens only. For comparison,
Table VIII shows the ratios and ranges for the adults of western
or typical Nearctic braueri, a California series, and eastern
braueri, plus those for the few available specimens of braueri
from central Europe. Third-instar larvae and puparia of western
and eastern braueri differ slightly as shown in figure 46.

Unfortunately, too few eastern adults were available in
measurable condition for the ratios to be strictly comparable with
those based on longer series of western examples. The influence
of size was also realized. Even so, the ranges of eastern and
western measures overlap more or less broadly, and many of the
mean ratios do not differ greatly. The male genitalia show no
differences, and the biology and ecology, with the characteristic
subcutaneous habit, are the same. The larval characters also
showed virtually no differences, except for the presence of
circumstigmatal folds in some--but not ail--western specimens,
and differences in hypostigmatal folds that are not regarded as
significant. Puparia of P. braueri are distinguished from most
other Nearctic Protocalliphora by the virtual absence of cuticular
spines, whether as spines or as tubercles. Spines that do occur are
minute as tubercles, giving the impression of fine sandpaper.
The occurrence of at least one sub-annulus per segment further
separates this species from all others. Some time after
measurements and calculations had been completed for the 25

136‘ Protocalliphora

males and 25 females from a variety of western localities, we
received an excellent series of 12 males, 13 females from
Columbia, Tuolumne Co., Calif., (David C. Rentz), reared from
subcutaneous larvae on a young black-headed grosebeak. As a
test of the ratios being recorded for braueri, this series of
probable siblings was also measured and ratios calculated. The
results have been included in Table VIII. In all males and in all
but one female, the individual ratios of the California specimens
fell within the ranges of the main western series; the exception
was a trifling difference of 0.003 in the minimum for the frons:
head ratio. The mean ratios were remarkably close, differing at
most by only 0.03. This essential agreement encourages us to
believe that the ratios will usually be useful criteria for species
recognition in this genus.

Males with brown calypteres, darker than usual or dark
because of discoloration, can be confused with those of P.
hesperia superficially, although the unique characters noted
above permit precise differentiation. Guberlet and Hotson (1940)
recorded maggots subcutaneously in house sparrows, and reared
a few specimens that were identified as "Hesperia." One of these,
a male (Seattle, Wash., USNM Collection), has been seen, and as
surmised from the biology, it proved to be typical braueri.
Perhaps this confusion is at the root of an error in Hall (1948).
Hall’s figure G, plate 21, labeled "hesperia" in the explanation on
p. 414, was apparently intended to be braueri (as hirudo), from
the reference on p. 194. Unfortunately, also, the figure does not
emphasize the distinctive apices of both surstylus and aedeagus.

Shannon and Dobroscky (1924) described the females of
"hirudo" as having the calypteres "darkened" or "smoky."
However, the type series shows that the females, except for the
allotype, were dirty and discolored and apparently influenced the
description. Actually the calypteres are lighter than in the males,
varying from whitish yellow, with outer margins yellow, to deep
orange-yellow. Peus (1960) was misled by the original
description into believing that true hirudo was not sexually
dichromatic in the color of the calypteres.

Females of braueri have been confused with aenea in
identifications because of the similarity of color. For example,
Shannon and Dobroscky (1924) recorded as "P. splendida var.
aenea" a broken specimen, of which only mesonotum and wings
remained, reared "from brain of a living fledgling of sparrow
kind" [USNM]. The mesonotum of this still extant fragment
shows distinct accessory notopleural bristles, the associated
puparium is characteristically that of braueri, and the myiasis is

P. brauveri — 137

typical of braueri, hence the specimen assuredly is braueri,
fragmentary though it is. As a further example, Hall (1948)
listed specimens of "aenea" reared from cliff swallow nests
(Kamloops, B.C.) and from a sparrow fledgling (Hamilton,
Mont.), but these all proved to be braueri. A male from the
Montana series was apparently the basis for Hall’s description of
the male of "aenea" (p. 185), and the measurements for the female
of "aenea" must have been based on specimens from these
misidentified series, in whole or in part.

We have concluded that the "Corvallis," Oregon specimen of
braueri identified by Melander as P. azurea and so recorded by
Cole and Lovett (1921) [Oregon State U.] is actually a Lents,
Oregon specimen like one now in the U.S. National Museum of
Natural History. Both bear the Accession No. 1191, under which
a file card in the Department of Entomology, Oregon State
University, records two maggots sent in by H. A. Darnell from
Lents, Oreg., July 15, 1914, from "the head of a young bird about
to fly." By July 24, both had pupated, and adults emerged July
27. The USNM specimen, which came from Melander’s
collection, bears correct data. The Oregon State specimen has the
same lot number but is labeled, apparently erroneously, as
Corvallis, July 14, and came to be published that way.

The specimens from Puget Sound, Wash. (Plath), which
formed the type series of P. hirudo var. cuprea, were earlier
included in the type series of Phormia metallica Townsend,
although then noted as a different form that might "perhaps
prove to be a good subspecies, or it may even be specifically
distinct" (Townsend 1919). However, two other specimens of the
same data are females near metallica or halli, probably the latter.
The labels do not make clear whether this species and braueri
came from the same nest, from a mixed infestation, or whether
different nests were involved.

Unconfirmed published records that can be accepted as
braueri with reasonable confidence because of the typical myiasis
are those by Bergtold (1927), who reared two adults of
"splendida" from larvae found in an abscess in a large swelling
below the right eye of a house finch at Denver, Colo., and by
Hall (1948), who recorded a male and a female of "hirudo" from
larvae found under the skin of nestling yellow warblers at Fort
Collins, Colo.

A European species was recorded as hirudo by Heinz (1954)
and Zumpt (1956), based on identification by Sabrosky.
However, Peus (1960), who compared a pair from Wyoming (det.
Sabrosky) with European material, believed that the European

138 Protocalliphora

species was distinct and new, and also that both belonged to a
new genus distinct from Protocalliphora. The new taxa were
published as Trypocalliphora and T. lindneri. He also raised
doubt whether hirudo sensu Sabrosky was the same as typical
hirudo Shannon and Dobroscky, noting for the former yellow
squamae (calypteres) in the females, brown in the males, whereas
hirudo was described as having "smoky" squamae in both sexes.
As already noted, however, the type series of hirudo was
somewhat discolored: females normally have yellowish or
whitish-yellow calypteres. Thus in reality the characters given
by Peus for hirudo sensu Sabrosky are typical for true hirudo.
Both hirudo and Peus’s /indneri are now considered synonyms of
P. braueri (Hendel) [Revision by Rognes 1985].

The characters of proportion given by Peus in his key, e.g.,
the width of the frons, have a range of variation, and the
characters of color not only vary somewhat but are greatly
affected by the condition and/or maturity of the individual
specimens. From the descriptive notes given by Peus for braueri,
we are inclined to suggest that the original reared specimens were
slightly teneral and hence paler. In reared series of Nearctic
braueri before us, we find examples from the same bird or nest,
and thus presumably siblings, in which calypteres in the males
range from whitish yellow (teneral) through deeper shades of
yellow and orange-brown to moderately dark brown (mature).
The extremely narrow frons recorded for braueri may also stem
at least in part from immature reared specimens. The frontal
Stripe is a weak area that is almost the last to reach the mature
stage and harden into normal position.

We have seen authentic /indneri Peus, having before us eight
examples, including a male and two females from Schlorer’s
rearing at Diersheim in southern Baden, in other words from the
same bird and locality as the holotype and probably siblings of it,
though not part of the type series. Ratios from measurements of
the few available specimens of Jindneri have been included in
Table VIII as "Europe" for comparison with the measurements of
Nearctic braueri. It is noteworthy that the ranges of the
individual ratios fall within the ranges of western (typical) hirudo
for most proportions. The exceptions are generally trifling.
Admittedly the sample of Jindneri is too small for mean ratios to
have any significance--and note that Peus’s sample of Nearctic
hirudo consisted of one male and one female--but bearing in
mind the variation observed in our much greater sample of
hirudo, we remained unable to separate European Jindneri from

P. brunneisquama =: 139

Nearctic hirudo. From our data we are happy to agree with
Rognes (1985) in synonymizing both under the still older braueri.

Protocalliphora (P.) brunneisquama, new species
(Fig. 28, Map 14)

Protocalliphora hesperia, in part.--James, 1955: 25 [Los Angeles
Co., Calif., July 4, 1950].

P. sp.--Horning and Barr, 1970: 73 [Idaho].

P. hesperia, in part.--Whitworth, 1976: 19, 27 [Utah, chipping
sparrow].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic dark blue to bluish purple; calypteres dark brown to
brownish; in male, surstylus long and narrow (Fig. 28) and cerci
exceptionally long and narrow.

Male. Head chiefly black in ground color, parafrontal and
parafacial dark gray to silvery gray microtomentose, that of
parafrontal thinning posteriorly to shining black prevertical area;
parafacial without changeable reflecting spot. Body chiefly
shining metallic blue to bluish purple, thinly gray
microtomentose, anterior slope of scutum gray microtomentose
but rest of scutum sparsely so, viewed from behind only weakly
demarcating the shining stripes; abdomen more shining than
thorax, brighter blue, tergites 2 and 3 gray microtomentose when
viewed from behind at a low angle, tergites 4 and 5 shining.
Calypteres dark brown to brownish.

Frons of moderate width, sometimes narrowing slightly
anterior to median ocellus, at narrowest 0.10 times head width
(0.07-0.13/5) and somewhat broader than ocellar span (1.26x;
1.00-1.375/5); parafrontal strongly narrowed posteriorly to
polished prevertical area, with one row of black hairs along
narrow part of parafrontal, row widening below to adjoin
parafacial rows, each parafacial with 3-4 irregular rows of
numerous black hairs; parafacial usually wider than frons (1.35x;
1.00-2.00/5), 1.70 times ocellar span (1.50-2.00/5), and slightly
narrower than vibrissal interval (0.88x; 0.82-0.90/4); ocellar
tubercle continued forward as a shining sublinear acuminate
preocellar area; cheek moderately broad, its height 2/5 maximum
length of eye (0.41x; 0.40-0.43/5).

Thorax without accessory notopleural bristles; tympanic
membrane and postalar wall with a few short and inconspicuous
hairs. Fore tibia usually with one posterior bristle (two on left

140 = Protocalliphora

tibia in male from Utah, and two on both fore tibiae in male
from New Mexico, but only one on other three males and all five
females).

Terminalia: cerci exceptionally long and narrow; surstylus
exceptionally long and narrow, curved, distal third narrower than
base (Fig. 28); aedeagus as usual for the genus (cf. Fig. 6a).

Female. Body color as in male, but scutum more heavily gray
microtomentose, as viewed from behind, and the shining stripes
thus more clearly demarcated than in male; parafrontal and
parafacial brownish yellow microtomentose, latter with a
changeable reflecting spot just below the frontal bristles;
preocellar area brownish microtomentose, more or less subshining
but not polished.

Frons at vertex 0.30 times head width (0.30-0.31/6) and 3.48
times ocellar span (3.44-4.12/6); frontal vitta with one to three
or four coarse bristlelike setae on each side, mesad of frontal
bristles; parafrontal broad throughout, though narrowing
somewhat dorsally; parafacial moderately broad, twice width of
a 3rd antennal segment, nearly half width of frons
(0.47x;0.44-0.48/6), 1.65 times ocellar span (1.455-1.94/6), and
0.92 times vibrissal interval (0.84-1/00/6); cheek height 0.44
times maximum length of eye (0.325-0.49/6), or 0.47x
(0.46-0.49/4) if two ’runts’ are disregarded [The runts made no
difference in other ratios, and the undersized females were
counted].

Apparent normal length of adults, 9-11 mm. Two females
from Utah are obviously undersized, 6 and 6.5 mm.

Immature Stages. Two puparia are known, but these are with
the undersized females and the details cannot be relied upon
except in a general way, such as the presence of good-sized
spines.

Type Series. Holotype male and allotype, Idaho: Craters of
the Moon National Monument, July 3 and 4 (male), 1965, Malaise
trap (D. S. Horning, Jr.). In the U.S. National Museum of
Natural History.

Additional paratypes. California: male, Forest Home, San
Bernardino Co., Sept. 21, 1936 (A. J. Basinger) [CAS]; female,
Monterey Co., 11 km N Lucia, July 8, 1963 (P. H. Arnaud, Jr.)
[CAS]; male, Tanbark Flat, Los Angeles Co., July 4, 1950 (W. O.
Marshall), published James 1955 as "hesperia" [U. Calif., Davis];
female, White Cloud Campground, Nevada Co., May 31, 1964 (P.
H. Arnaud, Jr.) [CAS]. Colorado: female, North Cheyenne
Canyon, Colorado Springs, June 17, 1926 (E. C. Van Dyke)
[CAS]. Montana: female, Fairy Lake, Gallatin Co., 9500 ft, Aug.

P. chrysorrhoea 141

18, 1983 (W. L. Downes, Jr.), on white-flowered umbellifer
[Downes Colln.]. New Mexico: male, White Mts., North Fork
Ruidoso, about 8300 ft, Aug. 12 (C. H. T. Townsend), on flowers
of Solidago trinervata [Identified by unknown person as
Calliphora viridescens] [Brit. Mus. (Nat. Hist.)]. Utah: male, two
females, Mantua, Box Elder Co., July 7, 1971 (T. L. Whitworth,
nest of chipping sparrow; 3 females, Box Elder Co.,
Mantua-Devil’s Gate, June-July, 1969 (T. L. Whitworth, Malaise
trap) [USNM, Whitworth Colln.]. Washington: female, Malaga,
Chelan Co., June 11-13, 1970, Malaise trap [Wash. State U.].

Specimens Examined. 5 males, 12 females.

Distribution. Western, known at present only from the states
of California, Colorado, Idaho, Montana, New Mexico, Utah, and
Washington. June 11-13 (Washington) to Sept. 21 (southern
California). The few records suggest that this is a species of
montane areas.

Hosts. Known only from chipping sparrow (Utah). Most
available specimens were hand-collected.

Ecology and Biology. Unknown. The size of most of the
specimens suggests that the species is usually a parasite of larger
birds than chipping sparrow.

Variation. The small sample makes it difficult to rely on the
ratios stated in the description. Variation in the posterior bristle
on the fore tibia was noted in the description.

Remarks. P. brunneisquama is especially easy to recognize.
It is one of the few species with dark brown to brownish
calypteres in both sexes, and it is the only one of these that has
narrow surstylus. The broad parafacial is useful for recognizing
females among those with brown calypteres. P. lata has
unusually broad parafacial, and on the other hand hesperia and
hesperioides have obviously narrow parafacials. P.
brunneisquama is intermediate between those extremes.

Etymology. The specific name, a noun in apposition, is
derived from the Middle Latin brunneus, brown, plus the Latin
squama, a scale.

Protocalliphora (P.) chrysorrhoea (Meigen)
(Figs. 6b, 11, 47, Map 6)

Musca chrysorrhoea Meigen, 1826: 60 [Austria].
Protocalliphora chrysorrhoea.--Hough, 1899: 66 [Placed in new
genus].

142 __Protocalliphora

P. azurea (Fallén).--Engel, 1920: 249-258 [From bank swallow
nests, Dachau, Germany; male terminalia figured].

P. chrysorrhoea.--Hennig, 1939: 363-4 [Possibly an ecological
race on bank swallow].

Apaulina sapphira Hall.--Sailer and Lienk, 1951: 209 [From "cliff
swallow," error for bank swallow].

P. azurea chrysorrhoea.--Zumpt, 1956: 95, 96 [Central Europe;
subspecies, restricted to nests of bank swallows].

P. chrysorrhoea.--Gregor and Povolny, 1959: 43-45 [Comparison
with azurea; colored figures of each species].

P. (Orneocalliphora) chrysorrhoea.--Peus, 1960: 208-211 [Central
Europe; neotype, female, Aachen, Germany, from Meigen
material in Munich].

P. chrysorrhoea.--Whitworth, 1976: 19,20ff [Utah]; Whitworth,
1977: 4933-B.

Diagnosis. Sexes concolorous, monochromatic, dark bluish to
bluish purple; calypteres white; tympanic membrane and postalar
wall with numerous long black setae; fore tibia with two posterior
bristles; frons of male of moderate width; surstylus of male
digitate in profile.

Male. Parafrontal and parafacial grayish to brownish gray
microtomentose, viewed from above the parafacial with a mottled
appearance of irregular light and dark crossbands, because of
numerous transverse ridges, and with numerous coarse black
hairs in about five irregular rows. Body dark bluish to bluish
purple, occasionally with dark greenish tint; thorax and abdomen
concolorous, shining, abdomen highly shining with thin gray
microtomentum visible only from behind at a low angle, scutum
and scutellum duller with two broad sublateral stripes of gray
microtomentum demarcating three shining stripes, although to
the unaided eye the scutum appears almost uniformly dull.
Calypteres whitish with light yellow rims.

Frons of moderate width, narrowing slightly anterior to
ocellar triangle, at marrowest 0.115 times head width
(0.095-0.135) and much broader than ocellar span (1.63x;
1.29-1.86); preocellar area shining but rugose, elongate triangular;
parafrontal narrowing above but clearly evident and
microtomentose to level of median ocellus; frontal bristles
decreasing posterodorsad and becoming mere hairs, both the
frontal bristles and a row of parafrontal hairs extending as far as
level of median ocellus; parafacial broad, twice as broad as 3rd
antennal segment, width 1.22 times that of frons (1.07-1.50), 1.99
times ocellar span (1.67-2.29), and virtually equal to vibrissal

P. chrysorrhoea 143

interval (1.00x; 0.87-1.07/19); cheek height 0.44 times (0.40-0.48)
maximum length of eye.

Thorax with scutum thickly covered with long, erect, fine
hairs, those on presutural area half or more as long as presutural
acrostical and dorsocentral bristles; each notopleuron sometimes
with an accessory notopleural bristle; postalar wall with a
conspicuous patch of black hairs; tympanic membrane with
numerous long black hairs, which are especially conspicuous.
Fore tibia with two posterior bristles.

Terminalia: cerci distally acuminate and parallel (cf. Fig. 5b);
surstylus in profile straight and digitate (Fig. 11), approximately
parallel sided and of medium width throughout, rounded apically;
aedeagus (Fig. 6b).

Female. Color as in male, monochromatic, but scutum
notably duller than in male because the stripes of gray
microtomentum are wider and more distinct throughout, the
median shining stripe especially narrowed. Frons_ broad,
widening anteriorly, at vertex about 3/10 width of head (0.29x;
0.26-0.31) and 3.64 times ocellar span (3.22-3.88); preocellar and
prevertical areas heavily brownish gray microtomentose, the
former subshining just anterior to median ocellus; frontal vitta
with several pairs of coarse, bristlelike, mesoclinate black hairs
and a number of short and inconspicuous hairs just mesad of
each row of frontal bristles; parafrontal and parafacial as in male,
brownish gray microtomentose, with numerous coarse black hairs;
parafacial about half width of frons (0.53x; 0.48-0.58), nearly
twice ocellar span (1.93x; 1.67-2.25), and slightly greater than
vibrissal interval (1.07x; 0.94-1.21/17); cheek height 0.47 times
(0.42-0.54) maximum length of eye. Mesonotal hairs and bristles
shorter than in male.

Apparent normal length of adults, 9.5-11.5 mm.

Third instar. Peritreme closed; cuticular spines numerous
between stigmatal plates and in hypostigmatal area. Prothoracic
fringe 29 wt (12-62)/ 21. Cephalopharyngeal skeleton 950
(870-990)/6 in length.

Puparium. Length 8-10 mm., breadth 4.1 mm. (3.8-4.4)/8;
medium-thick walled, dull, brown. Posterior region (Fig. 47a, b):
stigmatal plates 230 pw (220-250)/16 in diameter; distance between
buttons 690 pw (620-780)/8 and across stigmatal plates 1410 u
(1180-1485)/8; stigmatal ratio 0.49 (0.45-0.52)/8; stigmatal area
not outlined in large plaques; mesostigmatal area with numerous
spines and folds; hyperstigmatal spines 41 mw (14-58)/100;
hypostigmatal area with numerous, weak concentric folds;
hypostigmatal spines numerous, 18 (6-31)/100; circumstigmatal

144 —_ Protocalliphora

folds present, weak and concentric. Dorsal cuticle (Fig. 47c) with
numerous spines, 43 w (18-68)/100; cuticular ridges prominent.
Ventral cuticle (Fig. 47d): ventral band ratio 0.73 (0.65-0.81)/16;
spines of anterior patch 23 pm (12-56)/100; medial band
pronounced, spines 38 pu (23-56)/100; posterior band pronounced,
present on all segments; cuticular ridges on ventral surface either
pronounced or somewhat reduced.

Type. The species was described from Austria, apparently
based on a single female. It could not be found by Peus (1960),
who thereupon designated a neotype from Aachen, Germany,
that had been identified by Meigen himself.

Specimens Examined. 83 males, 76 females.

Distribution. Holarctic; Alaska to Quebec, south to Utah and
Montana, in bank swallow nests, probably much more widely
distributed than the records indicate.

Alaska [all USNM]. 12 males, 9 females, Ladd Field,
Fairbanks, Aug. 6 and 16, 1951, six bank swallow nests in colony
along Chena River; 3 males, 1 female, 60 mi E of Tok Junction,
July 14, 1948, bank swallow nest (R. I. Sailer) (publ. Sailer and
Lienk 1951 as Apaulina sapphira from "cliff swallow", error for
bank swallow, Sailer in litt.); 10 males, 9 females, Willow (N of
Anchorage), July 31, 1974, bank swallow nest (G. E. Haas).

Canada [all CNC]. Alberta: 3 males, 3 females, Elkwater (S of
Medicine Hat), July 15, 1969 (J. A. Shemanchuk), bank swallow
nest. British Columbia: 2 males, 1 female, 3 puparia, Atlin, 2200
ft, Aug. 6-7, 1955, barn swallow nest (H. J. Huckel); 3 males,
Summit Lake, 4200 ft, mile 392 on Alaska Highway, Aug. 19-21,
1959 (E. E. MacDougall); 4 males, 4 females, 6 puparia, Toad
River Lodge, 4500 ft, mile 422 on Alaska Highway, Aug. 15,
1959 (R. E. Leech), nest of violet-green swallow in old bank
swallow burrow. Quebec: male, Gatineau Co., Mesham Twnp.,
June 21, 1973 (D. M. Wood); male, Duncan Lake near Rupert (30
mi N of Ottawa), July 31-Aug. 4. 1971 (J. F. McAlpine); male,
female, 4 mi N of Eardley (above Ottawa), Aug. 25, 1971 (D. M.
Wood). Yukon Territory: male with puparium, Rampart House,
July 17, 1951 (J. E. H. Martin), bank swallow nest.

Contiguous United States. Montana: 3 males, 1 female,
Bigfork, nest July 20, 1917, larvae pupated July 21, adults Aug.
5 (C. H. Danforth), "external par[asite] on bank swallow"
[USNM]; male, female, Hamilton, Aug. 4, 1951 (L. E. Hughes),
bank swallow nest [Wash. State U.]. Utah: 2 males, 2 females,
Cache Co., Benson, Aug. 4, 1970 (T. L. Whitworth), barn swallow
nest [USNM]; 36 males, 44 females, most with puparia, Cache

P. chrysorrhoea 145

Co., Hyrum (Whitworth), 10 different bank swallow nests
[USNM, USU, Whitworth Colln.].

Two possible records of chrysorrhoea can be mentioned, but
there are no voucher specimens to check on the species. Lincoln
(1931) referred to a report from Crystal Bay, Minn., of bank
swallows with an abundance of "fat maggots protruding from
their nostrils." A similar record of numbers of large, blood-filled
maggots attached to nestling bank swallows, encircling their
necks and on top of their heads, near a beach in Oceana Co.,
Mich., was reported to us by George C. Steyskal. In both cases
the bank swallow parasites might well have been chrysorrhoea.

Basis of description of immature stages: Larva, 3rd instar (6
specimens): Tok, Alaska, bank swallow, July, 1948. Puparium (8
specimens): Tok, Alaska, bank swallow, July, 1948 (7); Bigfork,
Mont., bank swallow, July, 1917 (1).

Hosts. Chiefly bank swallow, also violet-green swallow
(rare), and isolated records from barn swallow. Whitworth (1976)
listed two nests of house sparrow and one of barn swallow
infested by chrysorrhoea, along with 95 infested nests of bank
swallow. The barn swallow record was verified (Benson, Utah),
but one of the house sparrow infestations proved to be sialia, and
perhaps the other was also. Uncommon records still do not
seriously affect the unquestioned predominance of chrysorrhoea
in bank swallow nests. European rearing records of chrysorrhoea
are likewise from bank swallows, there called sand martins (e.g.,
Engel 1920, Zumpt 1956, Peus 1960, Nuorteva 1960, Nuorteva
and Jarvinen 1961). The published record of chrysorrhoea from
"cliff swallow" (Sailer and Lienk 1951) was an error for bank
swallow, according to information from the late Dr. Sailer.

Ecology and Biology. Apparently restricted--with rare
exceptions--to cavity nests of the bank swallow, both in North
America and in Europe.

The record from the violet-green swallow was noteworthy
and was investigated further. According to information kindly
furnished by the collector, Robin E. Leech, the violet-green
swallow nest was in one group of about 15-20 nest holes from
15-30 inches deep in the vertical face of a roadside gravel pit,
about 5-7 feet above the floor of the pit and 3-4 feet below the
upper edge of the vertical face. The data indicate almost
certainly, according to the late Dr. A. Wetmore of the
Smithsonian Institution, that the holes were made by bank
swallows and subsequently utilized by the violet-green swallow,
which normally uses ready-made holes or crevices. This is an
interesting confirmation of Bennett’s belief in a ecological basis

146 ~~‘ Protocalliphora

for the distribution of Protocalliphora. Parasitism on bank
swallows exclusively had seemed an example of host specificity
by chrysorrhoea and therefore an exception to the ecological
theory. The parasitism of a different host suggests that the
apparent host specificity is the result of there being, usually, only
one host species in the bank swallow niche.

An interesting observation was provided by J. A.
Shemanchuk of the Research Station of Agriculture Canada at
Lethbridge, Alta. He reared chrysorrhoea at Elkwater, Alta., and
reported (letter to G. E. Shewell) that "Some of the adults came
from pupae found in the nests and some were reared from larvae
that were removed from under the skin and breast muscles of
nestlings."

Variation. A long series of 28 males, 39 females--runt
specimens not checked--from 10 different nests of bank swallow
at Hyrum, Utah, provided a fine opportunity for the evaluation
of variation. In the critical character of posterior bristles on the
fore tibia, all but three tibiae in the 67 specimens showed 2
bristles and these had 3: a right tibia on a female, a right on a
male and a left on another male, the two males in the same nest.

Most of the Alaska and Utah specimens are of the normal size
for the species, but many of the Hyrum, Utah specimens are
undersized, the smallest barely 6.5 mm in length. Otherwise they
are typical chrysorrhoea, and one can conclude that these were
underfed.

An accessory notopleural bristle was present in 55 of 148
specimens available for checking when this character was noted,
but usually on one side only. The species is so different from
braueri that the presence of an accessory notopleural bristle will
give no trouble.

The hindmost presutural intraalar bristle is absent in some
specimens, or present on one side only.

Remarks. Protocalliphora chrysorrhoea has long been
recognized as the parasite of bank swallows, sometimes recorded
as P. azurea (Fallén), sometimes as a subspecies of azurea, and
recently as a distinct species, no doubt aided by its apparent host
specificity.

Among the Nearctic species, chrysorrhoea and sapphira are
easily distinguished by a character that they share with most of
the Palearctic species: tympanic pit and postalar wall setose, and
front tibia with 2-3 posterior bristles, rarely only one. The frons
of chrysorrhoea is considerably narrower than that of sapphira,
and the surstylus is broader and less curved (cf. Figs. 11 and 26).

P. chrysorrhoea 147

Puparia of this species are similar to those of avium, hirundo
and sialia, but can be distinguished by the shorter prothoracic
fringe. Palearctic examples of puparia have also been studied
although our description is based entirely on Nearctic specimens.

Detailed comparison was made of Nearctic and Palearctic
material to determine whether we were dealing with a Holarctic
species on the Holarctic host, or whether the Nearctic population
was a new Species or subspecies. This comparison yielded no
differences that can be considered reliable, in the face of known
variability and the range of ratios. The Palearctic series is
unfortunately short, but significant in content. Eleven males and
ten females from Europe were available for comparison,
including four from Finland and 17 from Dachau, Germany (E.
O. Engel), identified as "azurea" by Engel and reared from the
nests of the "Uferschwalbe" (bank swallow). The Dachau
specimens were part of what Peus (1960) referred to as the classic
series Of chrysorrhoea and the basis of the study by Engel (1920).

Nearctic males have a slightly narrower frons than the
Palearctic, ranging from 0.095-0.135 times the head width, mean
0.115, compared with a range in available Palearctic males of
0.13-0.14, mean 0.137. Many Nearctic males are somewhat
teneral, and the frons is slightly compressed, so the frons: head
ratio is undoubtedly low. Allometric difference may also be a
factor; a small male from Finland shows a frons: head ratio of
0.12, like those of small Nearctic specimens.

No appreciable difference could be found in the male
terminalia. Minor chaetotactic differences were not reliable, and
apparently represent variation, possibly differing in degree in
different populations, but the present samples are much too small
to say. European males more often had an accessory bristle
between the two notopleurals. Nearctic specimens more often
had one or more fine hairs near the bristlelike infravertical
(below and behind each inner vertical bristle). In European
males the postocular setae were slightly shorter, and only slightly
longer towards the midline, whereas in Nearctic specimens the
setae appeared slender--a natural tendency in smaller
specimens!--and the dorsal six on each side were obviously
longer than the others, but there were exceptions on both sides.

In females, Nearctic specimens commonly have several pairs
of coarse bristlelike mesoclinate black setae on the frontal vitta,
mesad of the frontal bristles, whereas the few European females
available show fine hairs, short or of moderate length.

Puparia of Nearctic and Palearctic specimens are virtually
indistinguishable, although puparia of the latter have slightly

148 — Protocalliphora

shorter spines, even though the puparia are larger, and slightly
fewer cuticular ridges.

Many, although not all, of the Dachau specimens are larger
than available Nearctic specimens. Recent European revisions,
usually citing the Dachau material, give a length of 13-15 mm
for the species, with 14 as the average, whereas the range in
Nearctic material, admittedly somewhat teneral, is 9.5-11 mm.
A few Dachau specimens are the same size as the Nearctic, as
are the Finnish examples. Perhaps northern specimens of the
species tend to be smaller. Also the amount of available food or
the length of time the larvae had for feeding would affect the
size of the reared adults.

Protocalliphora (P.) cuprina (Hall)
(Fig. 16, 48, Map 7)

Protocalliphora metallica.--Neff, 1945: 73-6 [Calif.].

Apaulina cuprina Hail, 1948: 191 [Calif., Mont.; other specimens
partly aenea, partly metallica].

Protocalliphora cuprina.--James, 1955: 25 [Calif.]; Hall, 1965: 926
[Catalog; original records repeated].

P. sp. II.--Whitworth, 1976: 17ff.

Diagnosis. Dichromatic, body in male shining metallic dark
blue to bluish purple, in female heavily gray microtomentose and
dull bluish to bluish green, with tergite 5 of abdomen bright
cupreous to cupreous green; calypteres brownish yellow in male,
especially the rims, white in female; surstylus digitate,
approximately straight.

Male. Parafrontal and parafacial gray microtomentose; palpus
black tipped. Body shining, metallic dark blue to bluish purple,
thorax darker, bluish black, obscurely and thinly dark gray
microtomentose, shining stripes only weakly demarcated.
Abdomen thinly gray microtomentose on tergites 2-4, when
viewed from behind at a low angle. Calypteres brownish to
brownish yellow, especially on rims.

Frons narrow, narrowing anterior to ocellar tubercle, at
narrowest 0.06 times head width (0.05-0.07) and 0.89 times
ocellar span (0.71-1.18); preocellar area narrowly acuminate,
more or less rugose and shining or subshining, sometimes only a
ridgelike line; parafrontal strongly narrowed posteriorly,
prevertical area shining; frontal bristles weak posteriorly, ending
well in advance of median ocellus; parafacial with numerous

P.cuprina 149

black hairs in 3-4 irregular rows, equal to or only slightly wider
than breadth of 3rd antennal segment, 1.74 times width of frons
(1.31-2.00), 1.55 times ocellar span (1.38-1.83), and 0.72 times
vibrissal interval (0.615-0.83); cheek height 0.38 times maximum
length of eye (0.34-0.42).

Thorax without accessory notopleural bristles; tympanic
membrane and postalar wall usually bare; fore tibia typically with
one posterior bristle.

Terminalia: cerci distally narrowly acuminate and parallel (as
in Fig. 5b); with parallel-sided surstylus of moderate width,
digitate or only weakly curved (Fig. 16); aedeagus as usual for the
genus (as in Fig. 6a).

Female. Dark bluish black thorax, scutum heavily gray
microtomentose, two broad gray stripes demarcating a median
stripe and two shining lateral areas, the striping evident to
unaided eye, unlike the obscurity of male scutum; abdomen with
tergites 2-4 shining bluish green in direct view, but gray
microtomentose when viewed from behind at a low angle, tergites
2 and 3 especially dull; tergite 5 cupreous green, usually not as
decidedly cupreous as in metallica; calypteres white.

Frons at vertex about 1/4 head width (0.26x; 0.24-0.27) and
3.20 times ocellar span (2.86-3.67); preocellar area dull, evenly
gray microtomentose; parafacial moderately haired, in three
irregular rows, a changeable spot along upper margin, width of
parafacial 0.48 times width of frons (0.44-0.54), 1.54 times
ocellar span (1.36-1.77), and 0.79 times vibrissal interval
(0.69-0.92); height of cheek 0.40 times maximum length of eye
(0.37-0.44).

Apparent normal length of adults, 7.5-8.5 mm. Specimens as
small as five mm have been seen.

Third instar. No specimens examined, characters derived
from the puparium. Prothoracic fringe 60 [& (20-90)/60.
Cephalopharyngeal skeleton 890 tt (775-980)/9 in length.

Puparium. Length 6.7 mm (6-7.3)/6; breadth 3.4 mm
(2.6-3.8)/18; thin walled, dull, light brown. Posterior region (Fig.
48a, b); stigmatal plates 150 u (130-180)/36 in diameter; distance
between buttons 500 wu (400-550)/18 and across stigmatal plates
1000 w (875-1180)/18; stigmatal ratio 0.50 (0.46-0.54)/18;
stigmatal area not markedly outlined in cuticular plaques or folds;
mesostigmatal folds vary from weak and virtually absent to
moderately distinct; hyperstigmatal spines numerous, 28 [L
(15-40)/120; hypostigmatal area with numerous, weak, irregular
folds; hypostigmatal spines numerous, 18 pw (8-40)/120;
circumstigmatal folds absent (Fig. 48a), or if present, weak and

150 _—_— Protocalliphora

concentric (Fig. 48b). Dorsal cuticle (Fig. 48c) with numerous
spines posteriorly directed, 25 pw (13-40)/120; cuticular ridges
weak or absent. Ventral cuticle (Fig. 48d): ventral band ratio 0.63
(0.52-0.68)/14; spines of anterior patch 15 ps (8-24)/70; anterior
band pronounced; medial band narrow, spines 20 pw (16-30)/70;
posterior band pronounced.

Type Series. The species was described from a series of 23
males and 35 females, type locality Riverside, Calif., from four
species of birds. A few paratype males are actually hesperia, the
female from Mary’s Peak, Oregon is aenea, and the female from
Mt. Hood, Oregon is metallica. Holotype in the U.S. National
Museum of Natural History, and paratypes in that collection and
the California Academy of Sciences.

Specimens Examined. 265 males, 337 females.

Distribution. Western, British Columbia to Montana, south
to California, Utah and Wyoming.

Canada: British Columbia (all from nests): 90 males, 92
females, Haney (39 males, 27 females, yellow warbler), Kamloops
and Lac du Bois, Kamloops (western kingbird), Okanagan
Landing (yellow warbler), Qualicum (barn swallow), Vernon
(house finch).

United States (reared material from nests with one exception,
noted): California: 26 males, 37 females, Fresno Co. (nestling
willow flycatcher, yellow warbler), Le Grand (western kingbird,
also adults labeled "pupae dug from ground under dove’s nest"),
Mt. Lowe (Los Angeles Co.), Riverside ("goldfinch", house finch;
bulk of type series of cuprina), San Bernardino Co. (Camp
Baldy), and Nevada Co. (Truckee, 6000 ft). Idaho: 10 males, 14
females, all with puparia, Franklin Co. (Emigration Canyon;
western flycatcher). Montana: 10 males, 4 females, Corvallis
(barn swallow), Ravalli Co. ("wild canary"). Utah: 129 males, 182
females, almost all with puparia: Box Elder Co. (Mantua,
Empidonax flycatcher, and Mantua-Devil’s Gate, in Malaise
trap), Cache Co. (Benson, barn swallow, western kingbird),
Franklin Basin (Empidonax flycatcher, western wood-pewee),
Logan (eastern kingbird, yellow warbler). Washington: 7
females, Puget Sound (one labeled paratype of cuprina but not
published), Seattle ("western" robin, two as paratypes of cuprina),
Tacoma. Wyoming: female, Fremont Co. (Riverton at Wind River
at Highway 789).

Basis of description of immature stages: Puparium (18
specimens): Ravalli Co., Mont., American goldfinch, June, 1935
(7); Kamloops, B. C., Arkansas (i.e., western) kingbird, (4);
Cache Co., Utah, western wood-pewee, August, 1971 (7).

P.cuprina 151

Hosts. A dove, house finch, flycatchers (dusky, western, and
willow), American goldfinch, eastern and western kingbirds,
western wood-pewee, American robin, barn swallow, yellow
warbler. Whitworth (1976) recorded it also from Audubon’s [i.e.,
yellow-rumped] warbler.

Ecology and Biology. Whitworth (1976) noted that cuprina
was "virtually the only species found in warbler and flycatcher
nests."

Variation. In P. cuprina, there is typically one posterior
bristle on a fore tibia, but occasionally there are two bristles on
one side or the other, and sometimes on both sides. Of 575
specimens checked for this character, 433 or 75.3% have only one
posterior bristle on each fore tibia, and this is thus considered the
typical number. Only 50 or 8.6% had two on each front tibia,
but even this is frequent enough to cause difficulty in using the
key. When the 92 specimens with two on one side or the other
are added in, 142 or 24.7% show two bristles that might confuse
the user of the key. One male from Haney, B. C. has two bristles
on the left side and three on the right. In all specimens, the
lower posterior bristle is regularly present and outstanding;
others, if present, are usually smaller, not always outstanding,
and often close to the lower rather than well spaced from it as
usual in species with a regular number of two bristles.

Two odd variants: A female from LeGrand, Calif., has the
reclinate upper orbital bristles short and hairlike, rather than the
normal well developed bristles, and a female from Seattle, Wash.,
has an accessory notopleural bristle on the left side.

Some color variation also needs mention. In small and/or
very teneral individuals, the 5th tergite is only slightly yellowish
tinted, a faint yellowish green rather than a cupreous green.
Finally, in a series from Vernon, B.C., two females out of 16
have brownish yellow calypteres like the male.

Remarks. P. cuprina is in a complex of species with digitate
surstylus, but with the advantage in the general key of having a
female with cupreous green Sth tergite like the northern and
eastern P. metallica. Unlike the avium (eastern)-asiovora
(western) pairing, however, cuprina and metallica differ in the
male terminalia and fall into different species groups, metallica
having the surstylus narrow and curved, cuprina straight and
digitate or only slightly curved. In addition the two differ
greatly in the spination of the puparium: Fine sandpaper
appearance in metallica, distinct spines in cuprina. In the key to
males, cuprina passes to a small group of species with brownish
calypteres, and in the key to females the cupreous 5th tergite

152 Protocalliphora

associates it readily with the eastern bicolor and the chiefly
eastern metallica.

P. metallica of Plath (1919c) from "western" robin was
cuprina, at least in part, judging from the two females included
in the type series of cuprina, plus several others of Plath’s
material from Puget Sound, Wash.

The series from yellow warbler at Okanagan Landing, B.C.,
was a mixed infestation of spenceri and cuprina. Luckily both
sexes of the two species are easily distinguished, the males by
distinct surstyli, the females by the color of the 5th tergites.

Puparia of cuprina are separated with difficulty from those
of bicolor and spenceri. They can be separated from those of
bicolor on the basis of the shorter prothoracic fringe, but from
spenceri only by a general evaluation of all the characters.

Protocalliphora (P.) cuprina var. or new species

Female, Koehler, N. Mex., (W. R. Walton) [USNM]; female,
McKinley Co., N. Mex., 4 mi S Fort Wingate, 8000 ft, July 10,
1966 (P. H. Arnaud, Jr.) [USNM]; female, Mexico: Morelos,
Cuernavaca, July 9, 1961 (R. & K. Dreisbach) [Mich. State U.];
female, Mexico: Durango, 11 mi W Durango, 7000 ft, July 13,
1964 (J. F. McAlpine) [CNC]; male, Mexico: Durango, 3 mi E El
Salto, 8200 ft, July 1964 (J. F. McAlpine) [CNC].

The four females are close to cuprina and conceivably could
be variants of that species. Both tergites 4 and 5 are coppery
green and quite shining, although tergite 4 is finely
microtomentose when viewed from behind at a low angle. It is
also possible that they represent a new species, hitherto
unrecognized, but these isolated females, unassociated with
puparia and males, cannot be properly diagnosed and should not
be described. They are another indication of the paucity of
information on the Protocalliphora fauna of the Mexican plateau
(cf. Map 1). The lone male could be the male of the species, but
only rearing of a good series from one nest and host can give that
assurance. A female, Utah; Grand Co., La Sal Mts., Warner
Camp, 9500 ft, July 21, 1968 (W. J. Hanson), is not in good
condition but may belong here [Whitworth Colln.].

P. deceptor 153

Protocalliphora (P.) deceptor, new species
(Figs. 5b, 9, 49, Map 4)

Protocalliphora sp., probably new.--McAtee, 1929: 110 [Beil,
Md.].

Protocalliphora.--Thomas, 1936: 46 [Hocking, Lawrence, and
Jackson Counties, Ohio].

Protocalliphora avium.--Brimley, 1938: 371 [Raleigh, N.C.].

Protocalliphora n.sp.--Nolan, 1959: 355 [Bloomington, Ind.].

Protocalliphora n.sp.--Goddard and Lago, 1983: 482 [Starkville,
Miss. ].

Diagnosis. Sexes concolorous, monochromatic, dark blue to
bluish purple in ground color but mesonotum of female heavily
gray microtomentose; calypteres white; frons of both sexes
narrow; surstylus of male broadly rounded, as in avium.

Male. Head with parafrontal and parafacial brilliant silvery
microtomentose, latter without changeable spot; back of head
chiefly white haired, with one to two irregular rows of black
hairs behind row of postocular setae. Body dark blue to bluish
purple, or slightly bluish green, gray microtomentose, the
microtomentum on scutum demarcating three broad shining
stripes, abdomen appearing highly shining but microtomentum
especially heavy on tergites 2 to 4 when viewed from behind, the
5th tergite shining. Calypteres white.

Frons narrow, its width 0.07 times width of head
(0.05-0.08/22) and barely greater than ocellar span (1.01x;
0.77-1.25/22), narrowing from vertex 2/5 way to antennal bases,
then suddenly widening, at narrowest the parafrontals separated
by about width of one; preocellar area polished, elongate,
narrowly acuminate; parafrontal and parafacial smoothly and
finely microtomentose, the microtomentum thinning in
prevertical area, a single row of fine hairs beginning about
midway of parafrontal, becoming two rows anteriorly and finally
merging with the three to four irregular rows on parafacial; latter
obviously wider than frons (1.65x; 1.43-2.20/22), 1.67 times
ocellar span (1.38-1.83/22), and much narrower than vibrissal
interval (0.76x; 0.65-0.79/21); cheek narrow, in profile little over
1/4 eye height, its greatest breadth approximately 1/3 (0.35x;
0.31-0.40/22) greatest length of eye.

Scutal hairs less dense than in some species; no accessory
notopleural bristles; tympanic membrane usually bare, rarely with
one to a few fine and inconspicuous hairs; postalar wall bare or
with few short black hairs. Fore tibia with one posterior bristle.

154 Protocalliphora

Terminalia: cerci relatively short, acuminate, inner margins
parallel (Fig. 5b); surstylus longer than the cerci and greatly
broadened (Fig. 9), resembling that of avium and asiovora (cf.
Figs. 7-9); aedeagus usual for the genus (as in Fig. 6a).

Female. Parafrontal and parafacial bright and shining gray
microtomentose, faintly yellowish tinted, not silvery as in male.
Mesonotum much more heavily and extensively microtomentose
than in male, the microtomentum of scutum demarcating three
relatively narrow shining stripes. Abdomen as in male, much
more shining than thorax, but tergites 2 to 4 actually gray
microtomentose when viewed from behind at a low angle, tergite
5 shining.

Frons like that of braueri, exceptionally narrow for a female,
obviously half or less width of an eye, 0.23 times width of head
(0.20-0.25) and 3.01 times ocellar span (2.56-3.43), widening
gradually from vertex to opposite antennal bases; frontal vitta
with a few fine hairs toward sides, a small irregular preocellar
area often partly polished; parafrontal almost equibroad
throughout, about half width of frontal vitta, evenly
microtomentose, including prevertical area, one row of
parafrontal hairs, becoming two rows anteriorly; parafacial
sparsely haired, with two to three irregular rows, its width 0.52
times that of frons (0.44-0.56), 1.56 times ocellar span
(1.25-1.82), and 0.77 times vibrissal interval (0.64-0.92/21);
cheek unusually narrow, its breadth 0.34 times maximum ener
of eye (0.29-0.37).

Apparent normal length of adults, 7.5-8.5 mm.

Third instar. Peritreme open. Prothoracic fringe 55 pw
(25-90)/81. Cephalopharyngeal skeleton 773 wt (675-805)/10 in
length.

Puparium. Length 7.7 mm (7.2-8.8)/3; breadth 3.4 mm
(3.3-3.4)/3; thin-walled, shiny, light brown. Posterior region
(Fig. 49a-c): stigmatal plates 175 wu (145-195)/15 in diameter;
distance between buttons 530 pu (375-610)/8, and across stigmatal
plates 1180 w (1040-1290)/8; stigmatal ratio 0.45 (0.42-0.47)/8;
stigmatal area outlined in large plaques; mesostigmatal spines
present only as small tubercles; mesostigmatal plaques and/or
folds present; hyperstigmatal spines usually as tubercles, 6 bu
(2-14)/100; hypostigmatal area normally without folds but if
present, weak and irregular; hypostigmatal spines numerous, as
tubercles, 4 (2-8)/100; circumstigmatal folds absent. Dorsal
cuticle (Fig. 49d) with definite but relatively few spines, 12 pu
(4-20)/100; cuticular ridges weak. Ventral cuticle (Fig. 49e,f):
ventral band ratio 0.63 (0.55-0.66)/16; spines of anterior patch 11

P.deceptor 155

pt (4-20)/100; medial band pronounced, spines 12 pw (2-29)/100;
posterior band pronounced.

Type Series. Holotype male, allotype, and 8 male, 5 female
paratypes, Falls Church, Va., reared July 23, 1929, from [house]
wren’s nest (E. A. Chapin), each mounted with puparium. In the
U.S. National Museum of Natural History. Additional
paratypes:Connecticut: 11 males, 13 females, Stamford, Aug. 1-4,
1936, from nest of eastern phoebe (S. W. Bromley Colln.)
[USNM]. Delaware: 3 males, 3 females, Newark, July 10, 14
(male), 1967, reared from pupae in wood thrush nest (C.
Strehl)[U.Del., USNM]; male, 5 females, Newark, June 6, 1968,
from northern cardinal nest collected May 27 (F. M. Stiner, Jr.),
4 with puparia, and 8 males, 6 females, Newark, July 1 (5 males),
Aug. 10 (female), and 22 (3 males, 5 females), 1970 (Vivie E.
Davis), reared from wood thrush nests [U. Del., USNM]; male,
female, Greenwood, emerged Aug. 1, 1968, reared from nest of
common grackle (F. M. Stiner, Jr.) [Stiner Colln.]. Georgia:
female, Atlanta, July 16, 1935 (P. W. Fattig) [USNM]. Indiana:
2 males, Bloomington, June 9, 1954, nest of prairie warbler (Val
Nolan) [USNM]. Maryland: 2 females, Bell, reared from house
wren nest collected July 8, 1927 (W. L. McAtee) [USNM] (basis
for record of Protocalliphora sp. in McAtee 1929: 110); 23 males,
11 females, with puparia, Plum Point, Calvert Co., May 27, 1986,
pupae from Carolina chickadee nest in deserted bluebird nestbox
(J. H. Fales), flies emerged May 30-June 5[USNM]. Mississippi:
11 males, 2 females, Starkville, Oktibbeha Co., May 11, 1981, ex
pupae in nest of Carolina chickadee (G. Hurst) [U. Miss.,
USNM]. Missouri: female, Kirkwood, Sept. 9, 1952 (W. Downes)
[Iowa State U.]. North Carolina: male, 4 females, Raleigh, July
9, 1937, reared from nest of [eastern] phoebe collected July 7 (C.
S. Brimley) [N. C. Dept. Agr.] (basis for record of "avium" by
Brimley 1938: 371). Ohio: [Ohio State U. and USNM, except as
noted]: 13 males, 13 females, near Hanover, Licking Co., May
31-June 5, 1945, from American crow nest (S. E. Airesman, E.
S. Thomas); 4 males, 3 females, "Neotoma," Hocking Co., June
23, 1934, from larvae in nest of prairie warbler collected June 10,
1934 (E. S. Thomas) (also 2 males, 2 females, same data, at Amer.
Mus. Nat. Hist.); female, Washington Township, Lawrence Co.,
July 17, 1934, from larva in nest of worm-eating warbler, July
8, 1934 (C. F. Long, E. S. Thomas) (also one male at Amer. Mus.
Nat. Hist.); male, 2 females, Jackson Co., July 30 and August
(female), 1935 (E. S. Thomas), from pupae in nest of Carolina
wren (C. F. Long). South Carolina: 4 males, 6 females, Savannah
River Plant of Atomic Energy Commn., | mi from junction of

156 —- Protocalliphora

Savannah River and Upper Three Run’s Creek, May 11, 12,
1959, from nest of Carolina chickadee (W. A. Tarpley) [USNM
and Tarpley Colln.]. Tennessee: 8 males, 4 females, Nashville,
May 16, 1940, reared from northern mockingbird nest, including
8 mounted with puparia (Mrs. Amelia R. Laskey) [USNM]; 2
males, 2 females, Weakley Co., 1979 (T. David Pitts), from nest
of Carolina chickadee [USNM]. Virginia [all USNM]: 15 males,
16 females, Stafford Co., at mouth of Aquia Creek, NE of
Fredericksburg, late May 1969, from nest of Carolina chickadee
(E. T. McKnight); 7 males, 7 females, Stafford Co., near mouth
of Aquia Creek, June 8-17, 1970, from nest of Carolina
chickadee collected June 7 (E. T. McKnight); 19 males, 9
females, Stafford Co., Marlborough Point at mouth of Potomac
Creek, June 5, 1970, from puparia collected May 30 in nest of
Carolina chickadee.

Non-paratype Material: Ten males, 22 females, broken or
undersized specimens, some with same data as cited in the type
series, have not been considered as paratypes. Those recorded
because of additional localities or hosts (all USNM) are as
follows: Maryland: 3 males, 3 females, Patuxent Refuge, Bowie,
July 5, 1944; Tennessee: 4 larvae on which the larval description
is based, Nashville, June 25, 1947, from nest of great crested
flycatcher (Mrs. Amelia R. Laskey); Virginia: female, 4 mi W of
Luray, Blue Ridge Mts., July 30, 1956, from northern cardinal
nest collected July 21 (A. Wetmore).

Specimens Examined: 156 males, 142 females.

Distribution: Eastern United States, Missouri to Connecticut,
south to Georgia and Mississippi.

Basis of description of immature stages: Larva, 3rd instar (3
specimens): Nashville, Tenn., great crested flycatcher, June 25,
1947. Puparium (9 specimens, all from type series): Nashville,
Tenn., northern mockingbird, May, 1940 (2); Falls Church, Va.,
house wren, July, 1929 (7).

Hosts: Northern cardinal, Carolina chickadee, brown-headed
cowbird [probable; one of three fledglings from infested wood
thrush nest, Newark, Del. (V. E. Davis)], American crow, great
crested flycatcher, common grackle, northern mockingbird,
eastern phoebe, wood thrush, prairie and worm-eating warblers,
and house and Carolina wrens.

Ecology and Biology. The hosts thus far known appear to
indicate ubiquitous occurrence, especially in cavity nests.

Variation: Some aberrations were observed in the orbital
bristles. In females of the Newark, Del., series reared from nest
of northern cardinal, one female has only one pair of proclinate

P.deceptor 157

orbitals and no reclinate orbitals, and one female lacks the
reclinate orbital on one side. The first specimen, lacking the
reclinate orbitals altogether, would have appeared to be a female
of braueri but for the body color.

The series from Stafford Co., Va., reared June 8-17, 1970
from a nest of Carolina chickadee, provided the smallest
Protocalliphora that we have seen. The smallest female measured
only 4.5 mm, compared with others of 7.5-8 mm from the same
nest.

Remarks. The very narrow frons and brilliantly silvery
parafacials of the males suggest most of the 13 species with such
a narrow frons, but in its northeastern range only bicolor, braueri,
metallica, and shannoni are involved. Of these, bicolor has
tergite 5 cupreous in the males, braueri is much different with
yellowish to yellowish brown calypteres and accessory notopleural
bristles, and metallica, even though having a narrow frons, has
the frontal vitta much wider than a parafrontal. But even if the
differences in the narrow-fronted individuals are not considered
very distinct by an inexperienced person or one without
well-identified material for comparison, they will be astonished
to dissect the male terminalia and find the surstylus as broadly
rounded as in avium and asiovora (cf. Figs. 7-9). The cerci are,
however, narrowly acuminate and parallel distally, not at all like
those of avium and asiovora. This combination of narrow frons
and broad surstylus is unique in the genus, as far as known at
present.

Females are also unusual. Their narrow frons and broad gray
parafrontals at once suggest braueri over any of the other species,
but braueri lacks the reclinate orbital bristles and has accessory
notopleural bristles. Females of deceptor also are concolorous
with the males, metallic bluish or bluish purple, whereas braueri
is a dichromatic species and the females are aeneous green. If
immature stages are available, the numerous spines will easily
distinguish deceptor from braueri.

Puparia of deceptor are similar to those of metallica and
shannoni. They can be separated from shannoni by the presence
of tubercles, rather than spines, in the hyperstigmatal area, and
from metallica by the longer spines of the dorsal and ventral
cuticle.

In the May 1969 lot from Stafford Co., Va., from the nest of
a Carolina chickadee, 149 puparia were found in the nest, but
many were parasitized by Nasonia vitripennis. Numerous Nasonia
also appeared from puparia collected from the nest taken near the
mouth of Aquia Creek in June 1970.

158 Protocalliphora

Etymology. Both sexes are very deceptive in appearance,
hence the name deceptor, a noun in apposition, from the Latin
deceptor, a deceiver.

Protocalliphora fallisi, new species
(Figs. 24, 50)

Diagnosis. Sexes concolorous, monochromatic, shining
metallic bluish to bluish purple; calypteres white; frons of male
of medium width, preocellar area narrowly acuminate, shining or
somewhat gray microtomentose; surstylus narrow, curved.

Male. Head chiefly black in ground color, parafrontal and
parafacial silvery gray microtomentose. Body shining, metallic
bluish to bluish purple, scutum slightly gray microtomentose on
anterior slope and in two narrow stripes that set off a broad
shining (as viewed from behind) median stripe. Abdomen
shining in direct view, but thinly gray microtomentose when
viewed from behind at a low angle. Calypteres white.

Frons of moderate width, 0.11 times head width
(0.085-0.13/21) and 1.44 times ocellar span (1.14-1.67/21);
ocellar tubercle typically with four small bristles behind ocellar
bristles; preocellar area narrowly acuminate, often somewhat
shining, partly somewhat gray microtomentose; parafrontal
strongly narrowed above to polished prevertical area, sparsely
haired, one irregular row on lower part, row ending about
midway; parafacial often with changeable spot along upper
margin, and with few to numerous hairs in about three irregular
rows, width obviously greater than breadth of 3rd antennal
segment, 1.15 times width of frons (1.04-1.38/21), 1.66 times
ocellar span (1.57-1.83/21), and 0.86 times vibrissal interval
(0.77-1.00/21); height of cheek 0.42 times maximum length of
eye (0.39-0.44)/21); occiput with many white hairs, only 2-3
irregular rows of black hairs behind and below the row of
postocular setae.

Thorax occasionally with accessory notopleural bristles;
tympanic membrane and postalar wall with few short, pale, weak
and inconspicuous hairs. Fore tibia with one or two posterior
bristles on fore tibia.

Terminalia: cerci long and narrow to acute apices (cf. Fig.
5b); surstylus narrow, curved (Fig. 24); aedeagus as usual in the
genus (cf. Fig. 6a).

Female. Concolorous with male, but scutum somewhat more
gray microtomentose on anterior slope and the two scutal stripes.

P. fallisi 159

Frons 0.29 times width of head (0.265-0.31/20) and 3.54 times
ocellar span (3.14-4.00/20); preocellar area dull, finely and
evenly dark yellowish gray to brownish gray microtomentose, as
are parafrontal and parafacial; parafacial often with changeable
spot along upper margin, and sparsely to moderately haired, in
2-3 irregular rows, width obviously greater than breadth of 3rd
antennal segment, one half width of frons (0.50x; 0.46-0.58/20),
1.77 times ocellar span (1.60-2.10/20), and nearly equal to
vibrissal interval (0.97x; 0.92-1.04/18); height of cheek 0.45 times
maximum length of eye (0.42-0.48/20).

Apparent normal length of adults, 7.5-8.5 mm. (One
specimen, a net-caught male from Ogoki, Ontario, is 9 mm, and
five specimens are unusually small, 5-5.5 mm, undoubtedly
underfed).

Third instar. Peritreme closed; spines present between
stigmatal plates and in hypostigmatal area. Prothoracic fringe 50
bt (25-78)/87. Cephalopharyngeal skeleton 765 wb (690-885)/20
in length.

Puparium. Length 8.0 mm (7.2-8.8)/30; breadth 3.7 mm
(3.2-4.0)/30; thick-walled, dull, brown. Posterior region (Fig.
50a): stigmatal plates 155 w (120-190)/42 in diameter; distance
between buttons 595 yu (430-700)/20, and across stigmatal plates
1225 mw (900-1380)/20; stigmatal ratio 0.46 (0.44-0.52)/20;
stigmatal area outlined in plaques; mesostigmatal spines
numerous; mesostigmatal folds and plaques absent; hyperstigmatal
spines 16 pu (4-41)/100; hypostigmatal spines numerous, 9
(4-20)/100; circumstigmatal folds absent. Dorsal cuticle (Fig.
50b) with few spines, 16 ut (4-41)/100; cuticular ridges weak or
absent. Ventral cuticle (Fig. 50c): ventral band ratio 0.32
(0.20-0.38)/40; anterior patch indistinct, spines of anterior patch
10 2 (4-20)/100; medial band reduced, spines 10 pw (4-20)/100;
posterior band vestigial.

Type Series. Holotype male, allotype, and 54 paratypes (26
males, 28 females), Ontario, southern Algonquin Park, near Lake
Sasajewan, reared from nests of red-winged blackbird and
swamp sparrow in 1955; paratype male, Ogoki, Ont., Aug. 12,
1952 (J. B. Wallis) [Last, CNC]. Holotype and allotype in the
U.S. National Museum of Natural History, paratypes there and in
the Canadian National Collection. The holotype and allotype
were reared from the nest of red-winged blackbird 55-174 (G.
F. Bennett).

Non-paratype Material. Six specimens (male and five
females), including four reared from common yellowthroat nest

160 = Protocalliphora

in 1954, are small or in poor condition and are not included in
the type series.

Specimens Examined. 29 males, 34 females.

Distribution: Now known from two localities in Ontario.

Basis of description of immature stages: Larva, 3rd instar (6
specimens): Algonquin Park, Ont., white-throated sparrow, July,
1954. Puparium (30 specimens, from the type series): Algonquin
Park, Ont., red-winged blackbird, June 1955.

Hosts. Red-winged blackbird (predominant), swamp
sparrow, common yellowthroat; also, from puparia only (det.
Bennett), common_ grackle, white-throated sparrow,
rough-winged swallow, downy woodpecker.

Ecology and Biology. The few specimens thus far reared
were from nests taken in marshes.

Variation. Specimens may have either one or two posterior
bristles on the fore tibia, with the former slightly predominating
in the available material. Of the 64 specimens, 36 had one
posterior bristle on each fore tibia, 15 had two on each, and 13
had two on one tibia or the other.

Remarks. P. fallisi is nearest to P. tundrae in many respects,
and at one time the two were believed to be the same species.
However, the appearance of the male frons and _ the
hyperstigmatal spines, in particular, as well as some minor and
somewhat variable characters of occipital hairs and the bristles on
the ocellar tubercle, have led us to believe that the two are
distinct species.

Puparia of fallisi are separated with difficulty from those of
aenea. Generally, fallisi has a ventral band ratio smaller than
0.40, a shorter prothoracic fringe, and occurs in nests built in
swamps, along river banks, etc.

The species was mixed with metallica in the swamp sparrow
and two red-winged blackbird nests. Females of metallica were
easily recognized by the cupreous fifth tergite, but males of the
two species are much more difficult to separate in such a mixed
infestation.

Etymology. The specific name, in the genitive case, is
dedicated to Dr. A. Murray Fallis, respected protozoologist and
parasitologist, director of the doctorate study by Bennett, and
good friend.

Protocalliphora (P.) halli, new species
(Figs. 19, 51, Map 8)

Protocalliphora sp. 1V.--Whitworth, 1976: 19ff. [Utah].

P.halli 161

Diagnosis. Sexes concolorous, monochromatic, shining
metallic dark blue to bluish purple; calypteres brown in male,
white in female; surstylus digitate.

Male. Parafrontal and parafacial dark gray microtomentose,
sometimes faintly brownish or brownish yellow tinted; palpus
orange-yellow, black tipped. Body shining dark metallic blue to
bluish purple, scutum dark gray microtomentose but three
shining stripes only obscurely demarcated, as viewed from
behind; abdomen shining in direct view, bright gray
microtomentose on tergites 2-4, viewed from behind at a low
angle. Calypteres brown, paler but still brownish in teneral
specimens.

Frons narrow, 0.08 times head width (0.07-0.09) and 1.07
times ocellar span (0.86-1.50); preocellar area typically dull and
more or less triangular but occasionally rugose and acuminate,
and rarely linear and shining; frontal bristles decreasing in size
posterodorsad, the weak ones ending before level of median
ocellus; parafrontal narrowed above to shining prevertical area,
middle portion with one row of parafrontal hairs, widening to
several rows below at merger with parafacial hairs; parafacial
with numerous black hairs in about four irregular rows, width
obviously greater than breadth of 3rd antennal segment, 1.47
times width of frons (1.22-1.75), 1.57 times ocellar span
(1.36-1.83), and 0.75 times vibrissal interval (0.65-0.85); greatest
height of cheek 0.41 times maximum length of eye (0.38-0.43).

Thorax without accessory notopleural bristles and usually
without hairs on tympanic membrane and postalar wall; fore tibia
typically with one posterior bristle (cf. Variation).

Terminalia: cerci distally narrowed to parallel and acute
apical portions (cf. Fig. 5b); surstylus digitate and approximately
Straight, very weakly curved (Fig. 19); aedeagus as usual for the
genus (cf. Fig. 6a).

Female. Concolorous with male, but scutum more heavily
gray microtomentose, and demarcation of shining stripes usually
obvious to unaided eye, viewed from behind; parafacial with
changeable spot along upper margin; calypteres white.

Frons 0.26 times head width (0.23-0.29) and 3.24 times
ocellar span (2.86-3.86); preocellar area entirely dull, heavily
microtomentose; width of parafacial obviously greater than
breadth of 3rd antennal segment, 0.50 times width of frons
(0.44-0.55), 1.61 times ocellar span (1.50-1.71), and 0.81 times
vibrissal interval (0.73-0.92); greatest height of cheek 0.44 times
maximum length of eye (0.40-0.47).

162 Protocalliphora

Apparent normal length of adults, 7.5-8.5 mm.

Third instar. No specimens examined, characters derived
from puparium. Prothoracic fringe 55 pm (30-90)/25.
Cephalopharyngeal skeleton 900 pw (850-925)/4 in length.

Puparium. Length 7.5 mm (6.7-8.2)/5; breadth 3.3 mm
(3.2-3.6)/9; medium thin-walled, dull, brown. Posterior region
(Fig. 5la,b): stigmatal plates 170 uw (150-200)/18 in diameter;
distance between buttons 600 pu (450-700)/9, and across stigmatal
plates 1215 w (925-1425)/9; stigmatal ratio 0.49 (0.46-0.53)/9;
stigmatal area outlined in cuticular plaques and_ folds;
mesostigmatal area with a few small plaques and weak folds;
mesostigmatal spines scarce; hyperstigmatal spines numerous, 29
WW (15-50)/90; hypostigmatal area with weak, irregular folds;
hypostigmatal spines numerous, 18 pu (8-30)/90; circumstigmatal
folds weak, concentric. Dorsal cuticle (Fig. 51c) with numerous
spines, 28 mw (15-50)/90; cuticular ridges, if present, weak.
Ventral cuticle (Fig. 51d): ventral band ratio 0.49 (0.40-0.60)/17;
anterior band pronounced, anterior band not distinct, spines 17
Lt (8-25)/90; medial band narrow, thinned on the midline, spines
20 pw (8-40)/90; posterior band much reduced on anterior
segments, absent on the posterior ones.

Type Series. Holotype, male, allotype, and 29 paratypes (20
males, 9 females), Utah: Cache Co., Mendon, July 11, 1971 (7.
L. Whitworth), barn swallow nest 1155. Type in the USS.
National Museum of Natural History, paratypes there and in the
Utah State University and Whitworth Collections. Additional
paratypes: British Columbia: 135 males, 98 females, Heiron
Lease, Kamloops, Lac du Bois at Kamloops, Porcupine, and
Vancouver, various dates July-Aug., 1934, 1937, 1941, 1946 (G.
J. Spencer), nine barn swallow nests [UBC, USNM]; 78 males, 98
females, Qualicum, July 16-24, 1982 (N. K. Dawe and R. A.
Cannings), six barn swallow nests [UBC, USNM]. California: 20
males, 16 females, Napa Co., Howell Mt., 3 km NNE Angwin,
396 m, June 27-30, 1980 (Hugh B. Leech), nest of black phoebe
[CAS, USNM]; 44 males, 46 females, Riverside, June 1936 (A. J.
Basinger), phoebe nest 17 [CAS, USNM]. Utah: 153 males, 110
females, Cache Co., Benson, Logan, Mendon, Newton, and
Smithfield, various dates July 9 and Aug. 4, 1970, July 4, 11, 21,
24, Aug. 7, 12, 14, and Sept. 2, 1971, from 27 different nests of
barn swallows; also 2 males, Rich Co., Walton Canyon, July 22,
1970 (T. L. Whitworth), probably nest of pine grosbeak [USNM,
USU, and Whitworth Colln.]. Wyoming: 20 males, 6 females,
Lincoln Co., La Barge, July 3, 1970 (Whitworth), nest of Say’s

P.halli 163

phoebe, and 3 males, Lincoln Sage, Aug. 12, 1971, nest of barn
swallows [USNM, USU, and Whitworth Colln.].

Non-paratype Material: Numerous males and females from
lots cited under the type series; also 4 males, 4 females,
California: Siskiyou Co., Macdoel, July 21, 1948 (W. W. Wirth),
swallow nest under bridge [USNM].

Specimens Examined. 557 males, 481 females, a large
number with puparia.

Distribution: British Columbia, California, Utah, Wyoming.

Basis of description of immature stages: Puparium (9
specimens): Three, La Barge, Lincoln Co., Wyo., Say’s phoebe; 6,
Benson, Cache Co., Utah, barn swallow.

Hosts. Pine grosbeak (probable), black and Say’s phoebes,
barn swallow (predominant).

Ecology and Biology. The few hosts suggest that halli prefers
cup-nesting birds. The overwhelming proportion of records from
nests of barn swallow may or may not be significant for this
species. The ready accessibility and number of barn swallow
nests would undoubtedly tend to swell the numbers, as seen in
the work of Whitworth in Utah and Spencer, Dawe and Cannings
in British Columbia. For example, Whitworth (1976) recorded
that out of 392 nests of that bird species, 121 were infested by
Protocalliphora and of these 114 were infested by species IV, i.e.
halli as here described. Barn swallow is of course an excellent
host for Protocalliphora, and it is attacked by at least ten species
of the genus, according to our records, although some of these
may be only occasional or adventitious parasites on barn swallow.

Variation. Like the other species in the "hesperia complex,"
hesperia, hesperioides, and cuprina, variation was noted in the
posterior bristles on the fore tibia. Of the 703 specimens that
could be counted, 628 (89.3%) had only one bristle on each fore
tibia, 31 had two on each fore tibia, and 42 had two on one side
(right or left) and one on the other. Two specimens lacked a
bristle on one side.

A very few specimens showed a slight yellow tint on the 5th
tergite, and in isolated specimens this variation would cause
confusion with cuprina.

A most disturbing variation was in the appearance of the
preocellar area in males, which is usually short triangular and
gray microtomentose, but which may appear somewhat rugose or
granulated and narrowly acuminate, and even rarely a narrow
shining line. Yet in the males of the long reared series the
combination of brownish calypteres, black-tipped palpus, narrow
frons, and narrow parafacial is so consistent that we believe the

164 — Protocalliphora

differences in the preocellar area in this species must be accepted
as variation.

The calypteres of most males are not heavily browned, but
are usually more brownish yellow, a paleness that we have often
noticed in reared series when the specimens have not been
allowed to mature fully. Sometimes only the outer rims are really
brownish. In any case, they are obviously different from the
white calypteres of females, and from white calypteres of males
of species so characterized.

Remarks. For a discussion of the difficulties in the "hesperia
complex," see Remarks under hesperia. Briefly, all four species
have brownish calypteres in the male, also brownish in females
of hesperia and hesperioides, but halli and cuprina have white
calypteres in the female. Good series of all the species, and
especially of the last two, are available to back up the
conclusions. Males with brownish calypteres have in the past
usually been identified as hesperia.

Puparia of halli are easily confused with those of tundrae,
and to a lesser extent, those of aenea and fallisi. They are
separated from the latter two by weak circumstigmatal folds and
presence of some mesostigmatal plaques or folds. Puparia of
halli are separated with difficulty from those of tundrae, mainly
on the basis that the posterior band, although reduced, is present
on the anterior 5-6 segments of tundrae but absent on halli.

Etymology. The specific name, in the genitive case, is
dedicated to the late David G. Hall, Jr., whose 1948 book on "The
Blowflies of North America" marked for Protocalliphora a major
advance after the revision by Shannon and Dobroscky (1924).

Protocalliphora (P.) hesperia Shannon and Dobroscky
(Fig. 17, Map 8)

Phormia azurea (Fallén).--Plath, 1919c: 373-8, in part [Puget
Sound, Wash. ].

Protocalliphora splendida var. hesperia Shannon and Dobroscky,
1924: 249, 251-2 [B.C., Wash., the latter part of Plath’s
"azurea")|; Shannon, 1926: 128 [in key].

Apaulina hesperia.--Hall, 1948: 192 [In part].

Protocalliphora hes peria.--Hall, 1965: 926 [sw. Canada, nw. U.S.];
Whitworth, 1976: 27, 36 ff. [Utah, in part].

Diagnosis. Sexes concolorous, monochromatic, shining dark
blue to bluish purple; calypteres brownish or brownish yellow in
both sexes; surstylus digitate, not curved.

P.hesperia 165

Male. Parafrontal and parafacial silvery microtomentose.
Palpus orange-yellow, black at tip. Body metallic dark blue to
bluish purple; scutum gray microtomentose, two broad gray
stripes demarcating three broad shining stripes, as viewed from
behind; abdomen shining in direct view but gray microtomentose
when viewed from behind at a low angle, 2nd and 3rd tergites
decidedly gray, 4th less so. Calypteres brownish or brownish
yellow in both sexes.

Frons narrow, 0.08 times head width (0.08-0.09/24) and 1.15
times ocellar span (0.94-1.33/24); preocellar area almost always
gray and narrowly triangular, rarely somewhat shining; frontal
bristles decreasing rapidly in length posteriorly, the rows ending
well before median ocellus; parafrontal strongly narrowed,
prevertical area shining, mostly polished black; parafacial with
numerous black hairs in 3-4 irregular rows, width obviously
although not greatly wider than breadth of 3rd antennal segment,
1.37 times width of frons (1.20-1.57/24), 1.57 times ocellar span
(1.33-1.83/24), and 0.74 times vibrissal interval (0.67-0.79/21);
height of cheek 0.41 times maximum length of eye
(0.37-0.44/24).

Thorax lacking accessory notopleural bristles and hairs on
tympanic membrane and postalar wall, occasionally a few short,
pale and inconspicuous hairs on tympanum. Fore tibia typically
with one posterior bristle (cf. Variation).

Terminalia: cerci distally acuminate and parallel (as in Fig.
5b); surstylus digitate, very weakly curved, of moderate width
(Fig. 17); aedeagus as usual in the genus (as in Fig. 6a).

Female. Concolorous with male, but scutum not as dark,
brighter gray microtomentose; parafrontal, parafacial, and
preocellar area light yellowish gray microtomentose, parafacials
each with changeable spot along upper margin.

Frons about one-fourth head width (0.26x; 0.23-0.28) and
3.19 times ocellar span (2.75-3.57), preocellar area dull, evenly
microtomentose; parafacial as in male, width about half that of
frons (0.49x; 0.41-0.57), 1.57 times ocellar span (1.36-1.83), and
0.81 times vibrissal interval (0.73-0.92/21); height of cheek 0.43
times maximum length of eye (0.40-0.46).

Apparent normal length of adults, 8.5-9.5 mm.

Immature Stages. A few puparia of hesperia are known, but
some confusion in records makes it unwise to describe them at
this time. The cuticular spines are distinct, and the prothoracic
fringe is well developed, although not long.

Type Series. Shannon and Dobroscky described their variety
hesperia ina brief two-line description, saying only "Type.--Cat.

166 ~—‘Protocalliphora

no. 26860 U.S.N.M." On the following page they stated that "A
large series of this variety was reared from a number of species
of birds in regions of Seattle, Washington, summer 1918 (O. E.
Plath)" and from a male, five females from Ainsworth, B.C., July
11, 1903 (A. N. Caudell). The Seattle and Ainsworth specimens
constitute the type series of hesperia, with a type number stated,
but unfortunately the authors did not specify either the sex or the
type locality. Hall (1948) supplied this information, recording the
type as the male from Ainsworth, B.C. Arguably Hall’s action
could be considered lectotype designation if the original
publication were considered not to have fixed the type, but the
point need not be belabored; under either interpretation the result
is the same.

The collection in the U.S. National Museum of Natural
History now contains the holotype and the five females from
Ainsworth, B.C., one of the females marked allotype, the other
females paratypes, and also three males, two females,
hand-lettered "Puget Sd./Wash." "OE. Plath," with two of the
males and the two females also bearing the USNM red "paratype"
labels. These were not mentioned in the Type Catalog but they
could have been received later and labeled with the same number.
One female with the same labeling, including a red USNM
paratype label, is in the E. S. Thomas Collection now at Ohio
State University’s Department of Entomology, probably an
exchange received from Aldrich or Hall.

A larger series, 13 males and 11 females, in the Museum of
Comparative Zoology, bears handwritten labels "Puget Sound
Wash." (several are also labeled "Plath") but not in the same
handwriting as the USNM specimens. They have Shannon’s own
characteristic identification labels, and 3 males, 2 females also
bear red labels "paratype" which were obviously cut out of the
USNM label. It seems probable that these were received on loan
by Shannon and so labeled by him for return. Almost all of these
are not hesperia but the closely related species here described as
hesperioides. No host labels are on the specimens, nor on the
Plath material that is true hesperia, so it is impossible to sort out
the hosts listed by Plath (1919c). A mixture of species in
material reared from a variety of hosts is not unexpected.

Specimens Examined. 75 males, 83 females.

Distribution. Western, Alaska, British Columbia to southern
California, Utah, and possibly Colorado.

Alaska: 21 males, 26 females, 5.5 km SE Portage (S of
Anchorage, at base of Kenai Peninsula), reared June 19-21, 1978,

P. hesperia 167

from nest (? gray jay or thrush) collected June 10 (G. E. Haas)
[USNM].

Canada: British Columbia: male, 5 females, Ainsworth [type
series, USNM}]: male, Kamloops, July 31, 1937, emerged Aug. 19
in laboratory (G. J. Spencer) [UBC]; female, Shames, 18 mi SW
Terrace, June 23, 1960 (B. Heming), on flowers of Heracleum
lanatum [CNC]; female, Gagnon Road, 6 mi W Terrace, June 24,
1960 (C. H. Mann) [CNC].

Contiguous United States: California: 14 males, 18 females,
Riverside, May 1936 (A. J. Basinger), house finch, nest 19 [CAS,
USNM; 3 males were paratypes of cuprina}; female, Riverside,
May 4, 1934 (A. J. Basinger) [CAS]; male, Lake Co., Clear Lake,
July 29, 1948 (W. W. Wirth), swallow nest under bridge [USNM];
6 males, 9 females, Modoc Co., Adin Bridge, emerged July
22-27, 1978, barn swallow nest (C. Gold) [U. Calif., Albany];
female, Modoc Co., farmhouse by Adin, Aug. 6, 1978, barn
swallow nest (C. Gold) [U. Calif., Albany]; 4 males, 2 females,
Tulare Co., Farmersville, May 11, 1939 (T. H. G. Aitken),
swallow nest [CAS, USNM]. Oregon: male, Jackson Co., Ashland
Mt. above timberline, Aug. 5, 1952 (H. A. Scullen) [Oregon State
U.]. Utah (all reared T. L. Whitworth from nests of American
robin): female, Cache Co., Malibu Campground, July 17, 1970;
5 males, same locality and date; male, female, same locality and
date; two males, one female, Box Elder Co., Mantua-Devil’s
Gate, June 22, 1969; 3 males, one female, Mantua, July 14, 1970
[USNM, USU, Whitworth Colln.]. Washington: 4 males, 3
females, Puget Sound (in part, in type series) (O. E. Plath)
[USNM}]; 3 males, Puget Sound (O. E. Plath) [MCZ]; 8 males, 10
females, Seattle, July 1918 (O. E. Plath) [CAS]; female, Mt.
Rainier, Burroughs Mt., Aug. 21, 1934(A. L. Melander) [USNM].

A single female, Colorado: Clear Creek Co., Doolittle Ranch,
9800 ft on Mt. Evans, July 18, 1961 (W. R. M. Mason) [CNC]
may be hesperia, but it is far from the other known distribution.

Hosts: House finch, American robin, barn swallow. The
Alaska series was reared from a nest listed as "? gray jay or
thrush." Whitworth (1976) recorded hesperia from Cassin’s finch
(puparia only) and chipping sparrow, but the puparia have not
been found for checking, and the specimens from chipping
sparrow are herein described as brunneisquama N. sp.

Variation. One posterior bristle on the fore tibia appears to
be typical for hesperia, but there is enough variation to cause
some trouble. Of the 115 specimens that could be counted, 82
had just one bristle and 19 had two. Fourteen specimens had one
on one side and two on the other, either on right or left.

168 — Protocalliphora

Remarks. Four western species, hesperia, hesperioides, halli,
and cuprina, may be termed the "hesperia complex". All four
may occur in the same area, may attack the same or similar host
species, have puparia with distinct spines and generally similar
habitus, and have males with brownish calypteres, moderate to
narrow frons, and digitate surstylus. P. cuprina is more distinct
than the others because of the female with cupreous Sth tergite,
whereas the first three have all blue females, but males of cuprina
have been confused with the others.

Difficult as it is to separate the first three species, the
uniformity in fairly large samples persuades us to recognize three
species, but identifications of single specimens or of one sex only
will still be hazardous. Good reared series with both sexes
represented will be reassuring, especially in associating males
with brownish calypteres with a characteristic female. Females
of hesperia and hesperioides have brownish calypteres like the
males, but females of halli and cuprina have white calypteres.
(Warning: Very teneral females of hesperia and hesperioides have
pale calypteres, sometimes quite whitish, but still not the real
white of species that have truly white calypteres). In addition,
of course, cuprina has the cupreous Sth tergite that will separate
halli and cuprina easily. Females of hesperioides have the
preocellar area irregularly polished black, whereas in the other
three the preocellar area is dull, evenly and heavily brownish
gray microtomentose. Finally, in both sexes, palpi are entirely
yellowish to orange-yellow in hesperioides, but brownish- to
black-tipped in the other three. Males are very similar, but
hesperioides has a preocellar polished line or low ridge in
addition to the yellow palpi.

Because of the similarities, the name hesperia has been widely
misused. For example, hesperia of Guberlet and Hotson (1940)
proved to be braueri, and hesperia of James (1955) and
Whitworth (1976) was a mixture of hesperia, hesperioides, and
halli.

Protocalliphora (P.) hesperioides, new species
(Figs. 18, 52, Map 8))
Phormia azurea (Fallén).--Plath, 1919c: 373-8, in part [Puget

Sound, Wash. ].

Diagnosis. Sexes concolorous, monochromatic, shining dark
metallic blue to bluish purple; calypteres brown in both sexes;
palpus entirely orange-yellow; frons of male narrow with
polished line in preocellar area; surstylus digitate.

P. hesperioides 169

Male. Parafrontal and parafacial gray microtomentose, the
parafacial tinted with yellow; palpus entirely orange-yellow, not
black tipped. Body shining dark metallic blue to bluish purple,
scutum dark gray microtomentose, obscurely demarcating three
shining stripes, as viewed from behind under the microscope,
but appearing entirely dark to the unaided eye; abdomen highly
shining in direct view, but tergites 2 and 3 gray microtomentose
when viewed from behind at a low angle, 4th tergite slightly so.
Calypteres brown in both sexes.

Frons narrow, 0.06 times width of head (0.05-0.07) and 0.83
times ocellar span (0.69-1.00); preocellar area a polished line or
low ridge; frontal bristles decreasing rapidly in length
posterodorsad, ending just before median ocellus in several short
and weak hairs; parafrontal strongly narrowed above to more or
less polished prevertical area, on lower half with one row of hairs
becoming two or three rows just before merging with parafacial
hairs; parafacial rather densely beset with numerous black hairs
in about four irregular rows, width of parafacial only slightly
greater than breadth of 3rd antennal segment, 1.77 times width
of narrow frons (1.50-2.00), 1.47 times ocellar span (1.29-1.75),
and 0.68 times vibrissal interval (0.61-0.77); height of cheek 0.37
times maximum length of eye (0.31-0.41); occiput with about two
rows of black hairs behind row of postocular setae.

Thorax without accessory notopleural bristles and hairs on
tympanic membrane and postalar wall; fore tibia with one
posterior bristle; calypteres brown, often quite dark.

Terminalia: cerci distally narrowed to acute, parallel apical
sections (cf. Fig. 5b); surstylus digitate and approximately
Straight, slightly wider toward base (Fig. 18); aedeagus as usual
in the genus (cf. Fig. 6a).

Female. Concolorous with male, but scutum more heavily
gray microtomentose and more clearly demarcating shining
Stripes, as viewed from _ behind under’ microscope;
microtomentum of both parafrontal and parafacial yellowish
tinted, parafacial with changeable spot along upper margin.
Frons 1/4 head width (0.25x; 0.23-0.27) and 3.08 times ocellar
span (2.81-3.57); preocellar area with irregularly shaped, polished
black spot; width of parafacial 0.45 times width of frons
(0.40-0.50), 1.40 times ocellar span (1.25-1.64), and 0.74 times
vibrissal interval (0.64-0.85); height of cheek 0.40 times
maximum length of eye (0.37-0.45).

Apparent normal length of adults, 8.5-9.5 mm. One
undersized male was only 5.5 mm long.

170 ~~ Protocalliphora

Third instar. Peritreme closed; numerous spines present
between stigmatal plates and on hypostigmatal area. Prothoracic
fringe (in combination with measurements derived from puparia)
75 mw (55-110)/70. Cephalopharyngeal skeleton 920 w
(750-1020)/7 in length.

Puparium. Length 7.3 mm (7.1-7.5)/7; breadth 3.2 mm
(2.9-3.3)/7; medium thick-walled, dull, brown. Posterior region
(Fig. 52a,b): stigmatal plates 180 w (155-195)/14 in diameter;
distance between buttons 460 pu (425-500)/7, and across stigmatal
plates 975 ww (900-1050)/7; stigmatal ratio 0.48 (0.40-0.55)/7;
stigmatal area outlined in cuticular plaques and pronounced
ridges; mesostigmatal spines numerous; mesostigmatal folds and
plaques numerous and pronounced, bisecting stigmatal area;
hyperstigmatal spines numerous, 30 tt (18-50)/70; hypostigmatal
area rugose with numerous irregular folds; hypostigmatal spines
numerous, 20 , (8-45)/70; circumstigmatal folds weak,
concentric. Dorsal cuticle (Fig. 52c) with numerous spines
directed both posteriorly and anteriorly, 27 ju (18-40)/70;
cuticular ridges weak. Ventral cuticle (Fig. 52d): ventral band
ratio 0.75 (0.71-0.83)/14; ventral cuticle with cuticular ridges;
anterior band pronounced, spines of anterior patch not markedly
differentiated from surrounding spines, 18 wz (8-30)/70; medial
band pronounced, spines 25 pm (15-40)/70; posterior band
pronounced, present on all segments.

Type Series. Holotype male, allotype, and 44 paratypes (19
males, 25 females), Riverside, Calif., May 1936 (A. J. Basinger),
goldfinch nest 18. Type and allotype in the California Academy
of Sciences, paratypes there and in the U.S. National Museum of
Natural History. Additional paratypes: British Columbia: male,
Mt. Revelstoke, 5400 ft, July 7, 1952 (G. J. Spencer) [CNC].
California: 19 males, 14 females, Riverside, May 1936 (A. J.
Basinger), house finch nest 11 [CAS, USNM]; 10 males, 15
females, Riverside, May 1936 (Basinger), house finch nest 5
[CAS, USNM]; 5 males, 3 females, Anaheim, July 1936
(Basinger), house finch nest 23 [CAS]; male, Anaheim, May 12,
1935 (Basinger), bushtit nest [CAS]; 2 males,, 3 females, Blodgett
Forest, El Dorado Co., emerged July 23, 1979 (C. Gold), nest of
dusky flycatcher [U. Calif., Albany]; 3 males, 4 females, Poison
Meadow, Fresno Co., nest of yellow warbler, puparia July 18,
emerged July 24, 1988, conifer forest at 6500 ft. (B. E. Valentine,
S. Boland) [USNM]; female, Tanbark Flat, Los Angeles Co., July
7, 1950 (P. D. Hurd) [U. Calif., Berkeley; hesperia in James
1955]; female, Mill Valley, Marin Co., April 1965 (P. H. Arnaud,
Jr.) [CAS]; 3 males, one female, Woodside, San Mateo Co., Aug.

P. hesperioides 171

1, 1954 (E. Hunter), reared but host not stated [CAS]; male, San
Mateo Co., Corte de Madera Creek, vicinity of Portola, July 18,
1964 (P. H. Arnaud, Jr.) [CAS].

Non-paratype Material (undersized, teneral, or damaged): 5
males, Anaheim, and one female, Woodside, duplicates of type
material; 8 males, 12 females, Monterey Co., Hastings Natural
History Reservation, Calif., 1969 (N. A. M. Verbeek), nest of
western flycatcher [CAS]; 3 males, 2 females, Berkeley, Calif.
[MCZ]; female, Santa Paula, Calif., July 23 (Dean Burk) [U.
Calif., Berkeley]; 10 males, 11 females, Puget Sound, Wash.
(Plath) [MCZ; three bear "paratype" labels under Shannon’s
identification label as P. splendida hesperia].

Specimens Examined. 91 males, 95 females.

Distribution. British Columbia, California, Washington.

Basis of description of immature stages: Larva, 3rd instar (4
specimens): Woodside, San Mateo Co., Calif., host unknown,
July, 1954. Puparium (7 specimens): Monterey, Calif., western
flycatcher, 1969.

Hosts. Bushtit, house finch, goldfinch, dusky and western
flycatchers, yellow warbler.

Ecology and Biology. Nothing is known beyond the names
of a few hosts.

Variation. Although almost all specimens have entirely
orange-yellow palpi, two males of the small El Dorado County
series and one male of the Riverside Nest 5 series have
black-tipped palpi. One female of Nest 5 lacked the typical
polished black preocellar area. Both of these variations appear to
be the occasional aberrant occurrences that we have come to
expect. Another specimen of Nest 5 has a broad parafacial, and
this may be a Stray sialia; its width seems too great to be mere
variation.

As with other western species, the posterior bristles on the
fore tibia show some variation, in a character used in keys to
some extent and even considered a generic character by Hall, for
Apaulina. Of 123 specimens that could be counted, 92 had only
one posterior bristle on both fore tibiae, but 14 specimens had
two. Seventeen had two on right or left tibia but not on both.

Remarks. The four species of the so-called hesperia
complex, hesperia, hesperioides, cuprina, and halli, all have
brown calypteres in the male, male genitalia essentially the same,
with digitate surstylus, and distinctly spined puparia. Males have
commonly been identified as hesperia. P. hesperioides is
distinguished from the other three by the entirely orange-yellow

172 Protocalliphora

palpi, not black tipped except in the three aberrant specimens
already noted. Males are further recognized by the polished
black line in the preocellar area. Females have brown calypteres
like hesperia, but unlike hesperia the preocellar area has an
irregularly shaped, polished black spot. Females of the other two
species of the complex, cuprina and halli, have white calypteres
and dull preocellar area, in addition to which cuprina has the 5th
tergite cupreous in the female.

Puparia of hesperioides can be confused with those of
seminuda but the longer prothoracic fringe and larger ventral
band ratio should separate the two species.

Etymology. The specific name is compounded of hesperia,
with which the new species has been confused, and -oi/des, like,
from the Greek -eides, like.

Protocalliphora (P.) hirundo Shannon and Dobroscky
(Figs. 14, 38, 53, Map 9)

Protocalliphora splendida hirundo (subsp.) Shannon and
Dobroscky, 1924: 249 (as hirunda), 251, 252 [Wash.].

. Sp.--Stoner, 1926: 207-8 [lowa, bank swallow].

. Splendida hirundo.--Shannon, 1926: 128 [In key].

. Splendida.--Woods, 1926: 12-13 [Mass.].

. hirundo.--Hall, 1948: 196 [Wash., B. C.]; James, 1955: 25
[Calif.]; Boyes, 1961: 557, pl. I [Cytology]; Boyes and van
Brink 1965: 543 [Cytology]; Hall, 1965: 926 [Widespread];
Boyes and Shewell, 1975: 442, 446 [Cytology]; Whitworth,
1976: 20ff. [Utah]; Whitworth, 1977: 4933-B.

ajeas) ask as)

Diagnosis. Sexes concolorous, monochromatic, shining dark
metallic blue to bluish purple; calypteres white; preocellar area
in both sexes dull, heavily microtomentose; surstylus short, broad
on basal portion.

Male. Head black in ground color; parafrontal and parafacial
strikingly bright, brilliant silvery microtomentose, the parafrontal
distinctly microtomentose posteriorly at least to level of median
ocellus. Thorax shining, metallic dark blue to bluish purple,
dorsum somewhat gray microtomentose, the microtomentum
weakly demarcating three broad shining stripes but dorsum still
appearing shining and unstriped to the unaided eye. Abdomen
concolorous with thorax, but usually brighter blue and more
shining, tergites 2 to 4 gray microtomentose when viewed from

P.hirmdo 173

behind at a low angle. Calypteres opaque whitish, outer rims
tinged with pale yellow.

Frons (Fig. 38) slightly less than medium width, only slightly
narrowed anterior to ocellar tubercle, at narrowest 0.10 times
head width (0.09-0.11) and obviously wider than ocellar span
(1.37x; 1.21-1.58); frontal vitta about 1.5 times the width of a
parafrontal; preocellar area dull, heavily gray microtomentose;
parafrontal distinct throughout, with two irregular rows of fine
black hairs on lower part, decreasing above to one row and then
disappearing before end of frontal bristles; parafacial of medium
width, about twice as wide as 3rd antennal segment, wider than
frons (1.28x; 1.17-1.50), wider than ocellar span (1.76x;
1.50-1.91), and 0.89 times vibrissal interval (0.82-0.96), with 3-4
irregular rows of fine black hairs; cheek moderately broad, not
half maximum length of eye (0.43x; 0.40-0.45).

Thorax without accessory notopleural bristles and without
hairs on tympanic membrane, and with only an occasional seta or
two on postalar wall. Fore tibia with one posterior bristle.

Terminalia: cerci distally acuminate, parallel (as in Fig. 5b);
surstylus relatively short and broad, and characteristically
broader on basal two-fifths (Fig. 14); aedeagus as usual in the
genus (cf. Fig. 6a).

Female. Concolorous with male, but microtomentum of
frontal vitta, parafrontal, and parafacial dark gray, especially the
last, with light brownish yellow tinge, upper part of parafacial,
opposite lunule, usually with microtomentum arranged so as to
produce a dark brown spot viewed from above, changing to a
pale spot against dark background when viewed from below;
dorsum of thorax duller than in male, heavily gray
microtomentose, the three mesoscutal stripes more distinctly
demarcated than in male. Abdomen as in male.

Frons at vertex 0.28 times width of head (0.26-0.30) and 3.58
times ocellar span (3.00-4.33), widening slightly anteriorly;
frontal vitta toward the sides with a few short weak hairs;
preocellar area dull, densely and evenly microtomentose;
parafrontal broad throughout, evenly microtomentose, including
prevertical area; reclinate orbital bristles present; parafacial
broad, slightly over twice width of 3rd antennal segment, 0.54
times width of frons (0.48-0.58), 1.92 times ocellar span
(1.62-2.33), and approximately equal to vibrissal interval (1.00x;
0.93-1.15/24); cheek 0.44 times maximum length of eye
(0.41-0.47).

Apparent normal length of adults, 8.0-8.5 mm.

174 Protocalliphora

Second instar. Cephalopharyngeal skeleton 540 /2 in
length. Prothoracic fringe 56 mb (37-75)/25 in length, spines
numerous. Posterior region: peritreme closed, button distinct;
stigmatal plates 70 uw /4 in diameter; hyperstigmatal spines
numerous, 13 pw (6-21)/25; hypostigmatal spines numerous, 5
(2-8)/25. Dorsal cuticle with numerous spines, 11 p (8-18)/25.
Ventral cuticle: spines of anterior patch either single or two- or
three-pronged, 11 uz (6-14)/25; medial band pronounced, spines
12 pw (8-19)/25; posterior band present.

Third instar. Peritreme closed, spines present between
stigmatal plates and in hypostigmatal area. Prothoracic fringe 57
pt (21-103)/48. Cephalopharyngeal skeleton 850 uw (750-1000)/18
in length.

Puparium. Length 8 mm (7.2-8.9)/15; breadth 3.5 mm
(3.1-3.9)/15; medium thick-walled, dull, brown. Posterior region
(Fig. 53a, b): stigmatal plates 180 uw (150-200)/39 in diameter;
distance between buttons 550 wu (430-600)/20, and across
stigmatal plates 1230 jm (960-1410)/20; stigmatal ratio
0.46(0.42-0.48)/20; stigmatal area not outlined in plaques;
numerous mesostigmatal spines present; mesostigmatal plaques
absent, mesostigmatal folds absent or present, if present, only
two or three; hyperstigmatal spines numerous, 31 uw (21-41)/100;
hypostigmatal area with few, pronounced irregular folds;
hypostigmatal spines numerous, 14 uw (6-33)/100; circumstigmatal
folds pronounced, irregular, showing little variation. Dorsal
cuticle (Fig. 53c) with numerous spines, directed both posteriorly
and anteriorly, 31 w (21-41)/100; cuticular ridges pronounced,
irregular, few. Ventral cuticle (Fig. 53d): ventral band ratio 0.53
(0.40-0.74)/20; spines of anterior patch 17 (10-29)/100; medial
band pronounced, spines 27 pw (16-37)/100; posterior band
present on all segments, but weak.

Type Series. Holotype male and allotype, Stratford, Wash.,
July 4, 1920 (R. C. Shannon), from cliff swallow nests. Type
No. 26859 in the U. S. National Museum of Natural History.
Paratypes: 6 males, 2 females, same data as holotype [USNM], one
female, same data as holotype [MCZ]. The original description
listed the series as 35 males, 19 females, but the above are the
only specimens labeled as paratypes that have been found. One
male [MCZ], male, 4 females [USNM], one male, one female
from Belkin Collection [now in USNM], and 7 males, 3 females
from Melander Collection [now USNM] bear the same data but
are not labeled as paratypes, possibly an oversight. The ten from
the Melander Collection had been labeled P. chrysorrhoea,
possibly by Melander.

P.dinmdo 175

Specimens Examined. 270 males, 253 females.

Distribution. Northern transcontinental and western. Alaska
to Quebec, south to California, New Mexico, Iowa, and
Massachusetts. April 11 (Weld Co., Colo.) to Dec. 11 (San Luis
Obispo Co., Calif.).

Alaska. Male, female, Alaska Highway bridge over Tok
River, nest of cliff swallow; female, Chicken, under Taylor
Highway bridge, Chicken Creek, nest of cliff swallow; 4 males,
3 females, Tok Junction, nests of various Hirundinidae.

Canada. British Columbia: 47 males, 58 females, numerous
puparia, Atlin, Cariboo, Deadman’s Creek (near Walhachin),
Kamloops, Lac du Bois (near Kamloops), Minnie Lake Ranch
House (Nicola Ranges), Mission City, 108 Mile, Toad River
Lodge (Mile 422, Alaska Hgwy), and Vernon; best series, 23
males, 28 females, Kamloops, July 12-20, 1934 (G. J. Spencer),
from "eave" [i.e., cliff] swallow. Ontario: 56 males, 39 females,
numerous puparia, Algonquin Park, 1952-1956, chiefly barn and
bank swallow nests; 3 males, 4 females, Port Stanley. Quebec:
female, Knowlton. Yukon Territory: male with puparium,
Rampart House, nest bank swallow; male, Km 155 on Dempster
Highway.

Contiguous United States. California: 87 males, 75 females,
numerous puparia, localities in Del Norte, Kern, Los Angeles,
Modoc, Riverside, Sacramento, San Joaquin, San Luis Obispo,
Santa Barbara, Solano, Ventura, and Yolo Counties; best series,
28 males, 29 females, many puparia, Point Mugu, Ventura Co.,
June 23, 1949 (J. N. Belkin), from cliff swallow nest. Colorado:
male, 2 females, Larimer Co., cliff swallow nest; 8 males, 7
females, Weld Co., barn and cliff swallow nests; male, 2 females,
Weld Co., nest of yellow-headed blackbird. Idaho: 3 males, 2
females, Bonner’s Ferry, nest of small bluebirds; 2 males, 2
females, SW of Dickey, Custer Co., swallow nest. Iowa: 4 males,
3 females, 8 puparia, 40 larvae, Bluffton, Winneshiek Co., on
nestlings and in nest of cliff swallows. Massachusetts:
Huntington, male from barn swallow (Woods 1926 as P.
splendida), female from bluebird. Michigan: male, 5 females,
Midland. Montana: male, Granite Co.; 6 males, 11 females,
Hamilton, part from bank swallow nest; 6 males, 6 females,
Ravalli Co., part from swallow nests. New Mexico: 5 males, 5
females, 7 puparia, Rio Arriba Co., Cafones Creek, 2.5 km SW
Abique Dam, 6260 ft, Aug. 23, 1971 (N. Wilson), cliff swallow
nests [USNM, U. Northern Iowa]. New York: male, Syracuse,
bank swallow nest (P. splendida by Stoner 1936); 3 males, 2
females, Cranberry Lake, St. Lawrence Co., barn swallow.

176 = Protocalliphora

Oregon: male with puparium, Nestucca River, Tillamook Co.,
cliff swallow; 2 males, 1 female, Corvallis; 2 males, 3 females,
Chiloquin, Klamath Co., purple martin nest. South Dakota: 7
puparia, Redig, Harding Co., cliff swallow nest. Utah
(Whitworth): 4 males, 4 females, with puparia, Randolph, Rich
Co., cliff swallow nest; 9 males, 7 females, Franklin Basin, Cache
Co., tree swallow nest; 4 males, 6 females, Tony Grove, Cache
Co., purple martin nest. Washington: 2 females, Almota
(syntypes of sialia); 4 males, Stratford, cliff swallow nest.
Wyoming: male, Rock River.

Basis of description of immature stages: Larvae, 2nd instar
(2 specimens): Algonquin Park, Ont., bank swallow, July 1955.
Third instar (5 specimens): same data. Puparium (20 specimens):
Algonquin Park, Ont., barn swallow, August, 1955 (12);
Hamilton, Mont., bank swallow, date unknown (3); Ravalli Co.,
Mont., bank swallow, date unknown (3); Ventura, Calif.,
"swallow", June, 1949 (2).

Hosts: Yellow-headed blackbird, eastern bluebird, "a
bluebird" (Idaho), common grackle, purple martin, eastern
phoebe, European starling, and various swallows (bank, barn,
cliff, and American tree). Not seen, but recorded by Whitworth
(1976) from violet-green swallow. Commonest in swallow nests.

Ecology and Biology. The species appears to be commonest
in the West in cliff swallow nests. In Algonquin Park, Ont., it
was common in nests (e.g., barn swallow) in man-made
structures, and in cavity nests, particularly those in marshes: It
is one of several species, in addition to chrysorrhoea, that will
invade bank swallow nests.

Variation. In some hirundo the postocellar bristles (often
called postverticals) are well developed, being strong and
bristlelike, approximately equal in length and strength to the
ocellar bristles, and two to three times the length of the
postocular setae. In Protocalliphora in general, the postocellars
are not at all conspicuous, being less than or subequal to the
postocular row of setae. When this character was tabulated for
hirundo, we found that all western specimens had the short
postocellars typical of the genus. In Algonquin Park material,
however, over 2/3 of the specimens (both sexes) had strong
postocellars. Material from four nests had uniformly long
bristles, including one series of 50 specimens from a barn
swallow nest. Six nests (total 20 specimens) were uniform for
short postocellars. Three nests were mixed: 6 long, 4 short; |
long, | short; and 3 long, 6 short, respectively. The exact nature
of this character is unknown, but it is interpreted from present

P. interrupta’ 177

evidence as probably a Mendelian variant, perhaps a mutation
that has become or is becoming established in one part of the
range of hirundo.

Remarks. P. hirundo is one of a large group of species that
are usually somewhat difficult to distinguish. In males the short
broad surstylus (Fig. 14) is quite distinctive, and the combination
of silvery parafrontal and parafacial with the evenly
gray-microtomentose preocellar area makes the males of this
species among the easiest to recognize. Compared with sialia,
shannoni, and others near it, the habitus of the head is
outstanding. In the width of the frons and the silvery parafacial,
hirundo resembles sialia but that species has a_ polished,
acuminate preocellar area, as well as longer digitate parallel-sided
surstylus. Females of hirundo are less distinctive, but the broad
parafacial and entirely gray microtomentose preocellar area will
usually separate it readily from other species in the sialia
complex.

Puparia of hirundo are similar to those of avium, sialia and
chrysorrhoea. They can be generally separated by the fewer
circumstigmatal folds (which are quite characteristic of this
species) and their irregular pattern.

Protocalliphora (P.) interrupta, new species
(Figs. 30, 54, Map 7)

Protocalliphora sp. V.--Whitworth, 1976: I5ff. [Utah].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic bluish to bluish purple; calypteres white; surstylus long
and slender, strongly curved; puparium with minute spines,
appearance like fine sandpaper.

Male. Body shining metallic dark bluish to bluish purple,
scutum with gray microtomentum in two broad stripes that
demarcate three broad shining areas, as viewed from behind;
abdomen brighter than thorax, shining in direct view but tergites
2 and 3 distinctly and tergite 4 thinly gray microtomentose when
viewed from behind at a low angle. Calypteres white.

Frons narrow, at narrowest averaging 0.09 times head width
(0.075-0.11) and 1.18 times ocellar span (1.00-1.40); preocellar
area narrowly triangular, usually rugose and shining, rarely dull
gray microtomentose; parafrontal weakly haired on lower 1/3 to
2/5, narrowed above, narrowly microtomentose to or above level
of median ocellus; parafacial silvery gray microtomentose,

178 — Protocalliphora

sparsely haired in 2-3 irregular rows, width slightly greater than
breadth of 3rd antennal segment, 1.31 times width of frons
(1.00-1.50), 1.54 times ocellar span (1.23-1.80), and 0.70 times
vibrissal interval (0.53-0.80/23); height of cheek 0.41 times
maximum length of eye (0.36-0.415); occiput chiefly white
haired, only 1-2 irregular rows of black hairs behind and below
postocular setae.

Thorax without accessory notopleural bristles and with none
to few weak hairs on tympanic membrane and postalar wall.
Fore tibia usually with one posterior bristle.

Terminalia: cerci distally long and narrow to acute apices (cf.
Fig. 5b); surstylus long, exceptionally slender on _ distal
three-fifths, strongly curved (Fig. 30); aedeagus as usual in the
genus (as in Fig. 6a).

Female. Concolorous with male, but scutum somewhat more
heavily gray microtomentose, and shining stripes consequently
more clearly demarcated.

Frons of moderate width, at vertex 0.29 times head width
(0.26-0.31) and 3.40 times ocellar span (3.14-3.92); preocellar
area entirely dull, heavily microtomentose; microtomentum of
parafrontal and parafacial brownish gray or yellowish brown
tinted; parafrontal broad and gray up to vertex; parafacial
sparsely beset with scattered hairs, definite changeable spot
usually present along upper margin, width greater than breadth
of 3rd antennal segment, 0.46 times width of frons (0.43-0.52),
1.58 times ocellar span (1.43-1.83) and 0.78 times vibrissal
interval (0.69-0.85); cheek height 0.42 times maximum length of
eye (0.39-0.43).

Apparent normal length of adults, 7.5-8 mm.

Third instar. No specimens examined, characters derived
from the puparium. Prothoracic fringe 45 (24-80)/50.
Cephalopharyngeal skeleton 810 (750-925)/7 in length.

Puparium. Length 6.8 mm (6.8-6.8)/5; breadth 2.9 mm
(2.8-3.1)/7; medium thick-walled, shiny, light brown. Posterior
region (Fig. 54a): stigmatal plates 150 (150-150)/14 in diameter;
distance between buttons 560 (500-600)/7, and across the
stigmatal plates 1060 (950-1200)/7; stigmatal ratio 0.53
(0.46-0.57)/7; stigmatal plate faintly outlined in_ tubercles;
mesostigmatal spines few, represented primarily by small
tubercles; mesostigmatal folds absent, a few plaques present;
hypostigmatal spines scarce, small and many appearing as small
tubercles, 4 (3-5)/70; hypostigmatal area with a few plaques
but folds absent; circumstigmatal folds absent. Dorsal cuticle
(Fig. 54b) with few spines, 6 (4-8)/70; cuticular ridges

P. interrupta 179

weak or absent. Ventral cuticle (Fig. 54c, d): ventral band ratio
0.27 (0.16-0.38)/14; anterior band represented only by anterior
patch, which bears the largest spines of the puparium (except the
prothoracic fringe), 10 (8-12)/70; medial band present but
reduced, thinned on the midline and frequently interrupted;
spines small and usually represented as spine bases only, 4
(2-6)/70; posterior band vestigial or, more usually, absent.

Type Series. Holotype male, allotype, and 27 paratypes (10
males, 17 females), Utah: Cache Co., Benson, June 15, 1971 (T.
L. Whitworth), Nest 830, yellow-headed blackbird. Type,
allotype, and some paratypes in the U.S. National Museum of
Natural History, paratypes in Utah State University and
Whitworth Colln. Additional paratypes: Canada, British
Columbia: 8 males, 1 female, New Westminster, June 4, 1958
(Cox), nest of song sparrow; flies reared at Vancouver, June
4-20, 1958 by G. J. Spencer [UBC]. U.S.: Utah (all collected by
T. L. Whitworth, nests of yellow-headed blackbird except as
noted) [USNM, USU, and Whitworth Colln.]: 86 males, 70
females, Benson, June 18, 1970 and June 15 and July 4, 1971; 65
males, 65 females, Mendon, June 16, 1970 and July 11, 1971; 9
males, 7 females, Smithfield, June 5, 1971; 11 males, 12 females,
Wellsville, June 16, 1970 (1 male and 2 females from nest of
red-winged blackbird).

Non-paratype Material: Five males, five females from Utah
and a male from British Columbia have not been included
because of poor condition or unusually small size, but the data on
these are duplicated in the type seres. There is also a series of 2
males, 25 females, Bear River Refuge, Brigham City, Utah,
various dates April 29-June 26, 1965-1968, in trap (K. J.
Capelle) [Wash. State U.], which are undoubtedly interrupta, but
they are not reared and thus lack associated puparia.

A male from California: San Mateo Co., Corte de Madera
Creek near Portola, June 17, 1954 (P. H. Arnaud, Jr.) [CAS]
appears to be interrupta, but as a lone specimen, not reared, it is
not included in the type series. Three males, one female,
Ontario: Ogoki, Aug. 18, 1952 (J. B. Wallis), and one male,
Quebec: La Verendrye Provincial Park, Mi 61, Route 58, Aug.
20, 1965 (D. M. Wood) [CNC] appear to be interrupta, although
with some suggestion of spenceri. For either species, this would
be a great eastward extension of known range. Rearing will be
necessary to discover the puparial characters that would be
decisive.

Specimens Examined. 199 males, 203 females.

180 = Protocalliphora

Distribution: Utah (many specimens), British Columbia,
probably California, and possibly Ontario and Quebec.

Basis of description of immature stages: Puparium (7
specimens): Mendon, Utah, yellow-headed blackbird. All from
type series.

Hosts. Chiefly yellow-headed blackbird; also Brewer’s and
red-winged blackbirds, song sparrow, marsh wren (Specimens
from Brewer’s blackbird and the wren from Whitworth
identifications).

Ecology and Biology. Whitworth (1976) noted that this
species "was only found in marsh habitats where it commonly
infested nests of yellow-headed blackbirds." He reared it also,
but in small numbers, from Brewer’s and red-winged blackbirds
and the long-billed marsh wren. On the other hand, Spencer at
Vancouver, B.C., reared it from a song sparrow nest, so it may
also parasitize low-nesting birds that are not marsh-nesting.

Variation. Ten males and 2 females from Utah have two
posterior bristles on the fore tibia, and 6 males, 4 females have
2 bristles on one side, either left or right, but one on the other.

Remarks. The long series available makes it possible to say
with some assurance that this is a small species compared with
some others to which it comes close in the keys because of
various character states. Even in this small species, there are
occasional really small underfed specimens, the smallest only 5
and 5.5 mm in length.

The most remarkable feature is the minutely spined
puparium, which resembles that of metallica, a dichromatic and
chiefly eastern species. In the general key, the fact that males
and females of interrupta are concolorous takes it far from
metallica, but in the key to males the similarity of male surstyli
brings the two closer together. In some respects, males of
interrupta key out near parorum, but that species is consistently
much larger and differs in other respects as well, besides being
a parasite of chickadees. Females of interrupta are not distinctive
and end up with several other species near the end of the key.

The puparium differs from that of metallica in having the
hindmost of the usual three bands of spines lacking on the
ventral surface of the anterior end. Thus the absence of the third
band results in a strongly banded appearance of alternating spine
bands and bare areas. This is especially noticeable on about three
segments posterior to the cap. These segments have the full three
bands of spines in metallica. The medial band is present in
interrupta but thin and often interrupted on the midline.

P.lata 181

Puparia of interrupta can be confused with those of aenea,
metallica, spenceri, and tundrae. However, the small spines, lack
of cuticular ridges and the interrupted medial band serve to
readily distinguish it. The "sandpaper-like" puparium also helps
to distinguish interrupta from other species, such as fallisi,
parorum, and spenceri, with narrow curved surstylus and virtually
indistinguishable females.

Etymology. The specific name comes directly from the Latin
participle interrupta, referring to the break in the usual bands.

Protocalliphora (P.) lata, new species
(Figs. 10, 33, 55, Map 10)

Protocalliphora"gigantia" Whitworth, 1976: 17, 18, 27, 35, 37 (as
"gigantica") [Nomina nuda; Utah, especially on
Falconiformes].

Diagnosis. Sexes concolorous, monochromatic, shining dark
metallic blue to bluish purple; calypteres dark brown in both
sexes; both sexes dichoptic, frons as broad as an eye; surstylus
digitate but relatively short and therefore appearing rather broad
(Fig. 10); unusually large species, normal length 17-19 mm, the
largest Nearctic species of the genus.

Male. Head black in ground color; parafrontal and parafacial
yellowish to brownish yellow microtomentose. Body shining,
dark metallic blue to bluish purple; scutum thinly dark gray
microtomentose, barely interrupting the shine, the usual
mesoscutal stripes indistinct. Abdomen more highly shining than
thorax, with only a thin grayish "bloom" (microtomentum) visible
from behind viewed at a low angle. Calypteres dark brown to
brownish black.

Head unusual in genus in having broad frons (Fig. 33), as in
female; frons approximately as wide as an eye, 1/3 width of head
(0.34x; 0.32-0.35/8) and over four times as wide as ocellar span
(4.15x; 3.74-4.60/8); frontal vitta broad, with some bristlelike
setae along each side, just mesad of frontal rows of bristles; large,
polished black preocellar area; parafrontal broad, anieriorly as
broad as_ parafacial, narrowing strongly posteriorly,
microtomentose anteriorly but narrowed prevertical area shining
black; one or two pairs of proclinate orbital bristles and a pair of
reclinate to lateroclinate upper orbitals; parafacial extraordinarily
broad, width over three times that of 3rd antennal segment,
nearly 2/3 as broad as frons (0.645x; 0.595-0.66/8), 2.67 times

182 — Protocalliphora

ocellar span (2.27-3.00/8), and 1.83 times vibrissal interval
(1.47-2.00/8); cheek broad, 0.635 times maximum length of eye
(0.61-0.67/8).

Thorax without accessory notopleural bristles; tympanic
membrane nearly bare, with only two or three weak hairs;
postalar wall centrally with only two or three fine hairs,
occasionally up to six. Fore tibia with two posterior bristles;
claws strong, abruptly curved at right angles about midway.

Terminalia: cerci short, stout, straight (as in Fig. 5b);
surstylus moderately broad, wider at base and broadly rounded
distally (Fig. 10), the breadth making it seem intermediate
between the long digitate surstylus of sialia and the very broad
surstylus of avium; aedeagus as usual for the genus (as in Fig. 6a).

Female. Color as in male. Frons broad, at vertex 0.35 times
width of head (0.32-0.38/20), and 4.63 times ocellar span
(4.30-5.20/20), strongly widening anteriorly and projecting well
anterior to eyes; preocellar area large, polished black; frontal
vitta with a few fine hairs and some bristlelike setae on each side
mesad of frontal bristles; parafrontal broad, with numerous
coarse black hairs in from four to five irregular rows opposite the
lowest frontal bristles up to two rows at the reclinate orbital
bristles; prevertical area chiefly polished black; two proclinate
and one reclinate orbitals; parafacial very broad, with numerous
black hairs in about six irregular rows, its width 0.61 times width
of frons (0.57-0.66/20), 2.84 times ocellar span (2.60-3.12/20)
and 1.90 times vibrissal interval (1.53-2.23/20), the interval
appearing unusually narrow for Protocalliphora because of the
wide parafacial; cheek broad, its height 0.62 times maximum
length of eye (0.60-0.66/20).

Thorax as in male. Fore tibia likewise with two posterior
bristles.

Apparent normal length of adults, 11-12 mm.

Third instar. Peritreme closed, spines present between
stigmatal plates, numerous in hyperstigmatal area. Prothoracic
fringe 74 w (31-130)/30. Cephalopharyngeal skeleton 1085 wu
(880-1275)/11 (some measurements derived from puparia).

Puparium. Length 13 mm (on one); breadth 5.1 mm
(4.7-5.5)/10; thick-walled, dull, black-brown. Posterior region
(Fig. 55a, b): stigmatal plates 285 wu (250-325)/24; distance
between buttons 745 wu (425-1075)/12, and distance across
stigmatal plates 1700 mw (1250-2125)/12; stigmatal ratio 0.44
(0.34-0.51)/12; stigmatal area outlined in pronounced cuticular
plaques; mesostigmatal spines absent or very _ scarce;
mesostigmatal folds absent; hyperstigmatal spines pronounced, 37

P.lata 183

Lt (24-64)/120; hypostigmatal area with few, weak spines, 24 wu
(16-48)/120; hypostigmatal folds absent or at best, weak;
circumstigmatal folds pronounced, irregular. Dorsal cuticle (Fig.
55c) with numerous large spines, 45 w (24-64)/120; cuticular
ridges prominent, usually with two rows of spines associated with
each ridge, one directed anteriorly and the other posteriorly.
Ventral cuticle (Fig. 55d): ventral band ratio 0.63 (0.55-0.73)/25;
spines of anterior patch slender, 24 wu (8-40)/120; medial band
present, spines 40 pu (16-64)/120; posterior band reduced to one
to three rows of small spines.

Type Series. Holotype male, allotype, and 6 male, 7 female
paratypes, Utah: Smithfield, Cache Co., June 28, 1970 (T. L.
Whitworth; nest 208), nest of American kestrel [sparrow hawk].
Type in the U.S. National Museum of Natural History.
Additional paratypes: Utah [USNM, USU, Whitworth Colln.]: 2
males, 11 females, Mendon, Cache Co., July 20, 1970 (T. L.
Whitworth, nest 380), nest of American kestrel; male, Bunch
Grass Creek, Cache Co., July 6, 1970 (T. L. Whitworth, nest 340),
nest of red-tailed hawk. British Columbia: male, Vavenby, Mar.
26, 1934 (V. K. Moilliet) [Brit. Mus. (Nat. Hist.)]. Oregon:
female, Mt. Hood, 3000-6000 ft, June 26, 1925 (E. C. Van
Dyke) [CAS]. California: female, Summit Camp, Lassen Co.,
June 28, 1949 (E. L. Atkinson) [U. Calif., Berkeley]. Almost all
the Utah specimens were accompanied by puparia.

Non-paratype Material. 4 males, 7 females, Idaho:
Emigration Canyon, Franklin Co., June 28, 1970, nest of
red-shafted flicker [USU]. These specimens seemed so much
smaller than the "giant" ones that the possibility of still another
species had to be considered. The same fundamental
features--male terminalia, broad frons and parafacial in both
sexes, male with orbital bristles--were present although some
proportions differed, as often occurs in smaller specimens. We
finally concluded that these were conspecific with the type series,
merely undersized specimens, probably underfed, a not
uncommon occurrence in Protocalliphora. Also a headless male,
Logan, Utah, July 7, 1951 (collector unknown) [USU]. Also four
empty puparia, so large and heavily ridged that they are certainly
the puparia of Jata: Ravalli Co., Mont., Aug. 14, 1966 (Wm. L.
Jellison), from old nest of American kestrel [USNM].

Specimens Examined. 16 males, 28 females.

Distribution. Western, known at present from British
Columbia, California, Idaho, Oregon, and Utah, and probably
Montana (puparia only).

184 — Protocalliphora

Basis of description of immature stages: Larva, 3rd instar (3
specimens): Pasadena, Calif., brown [California] towhee, June,
1952. Puparium (10 specimens): Smithfield, Cache Co., Utah,
American kestrel (4); Mendon, Cache Co., Utah, same host (6).

Hosts. Red-shafted flicker, red-tailed hawk, American
kestrel [sparrow hawk], brown towhee. It has also been recorded
from the northern goshawk by Whitworth (1976).

Ecology and Biology. Unknown, other than the names of a
few hosts, which suggest--as does its size-- that the species
primarily attacks large birds. Whitworth (1976) suggested that it
attacked especially Falconiformes, which seems likely although
at this stage the records are few.

Variation. The size difference has already been noted, but
this is of common occurrence in Protocalliphora. Whitworth
reported (personal communication) that the small specimens
"were removed from a flicker nest as first and second instars and
reared on a young Brewer’s blackbird." Most specimens of this
series are 7-7.5 mm. in length, far below the normal length of the
species. Two males were 9 mm., still smaller than the normal
length.

Proclinate orbital bristles are present on one or both sides in
all males, but the numbers vary: 2 left and 2 right in seven males,
2-1 or 1-2 in three males, 1-1 in three, and 1-0 in one. Females
show the full two pairs in 27 of the 28 available, but only one
pair in the female from Mt. Hood, Oregon. Two posterior
bristles on each fore tibia is typical of the species, but there is
only one bristle on either left or right tibia in 2 of 15 males and
5 of 28 females.

Remarks. The headless male from Logan, Utah, was the first
one seen, and it was recognized at once as the largest Nearctic
Protocalliphora seen up to that time, with male terminalia that
also marked it as a distinct species. At the same time, two
slightly shorter females from Oregon and California were being
recognized as a distinct species because of their unusually wide
parafacials. Later, when Whitworth reared the type series of /ata
in Utah, the headless male and the two females were at once
recognized as /ata, and happily the species could be recorded
from a more extended range.

The structure of the head is unique among the known
Nearctic species, with the unusually broad parafacials, frons
broad in both sexes, and males with proclinate and reclinate
orbital bristles so that they appear at first glance to be females.
The dark brown calypteres in both sexes and the two posterior
bristles on the fore tibia, the latter coupled with the virtual

P. metallica 185

absence of hairs on the tympanic membrane and postalar wall,
also distinguish it from other species.

The specimens in the type series are by far the largest of the
known Nearctic species of Protocalliphora. The unusual size and
general appearance remind one at once of the large northern blow
fly Cynomya mortuorum (L.), which is a calliphorine and not a
phormiine calliphorid.

Puparia of /ata are separated from those of avium, hirundo,
and sialia, which they broadly resemble, by the generally larger
size (/ata is the largest of the Nearctic Protocalliphora). This
species is separated from both avium and hirundo by the generally
larger cuticular spines and the longer prothoracic fringe. It is
separated from sialia, which lata closely resembles in most
respects, by the fact that the prothoracic fringe of sialia is nearly
twice as long. All four species have marked cuticular ridges with
two rows of spines associated with each ridge, one row pointed
posteriorly, the other anteriorly.

Etymology. The specific name is derived from the Latin
latus, broad, referring to the unusually broad frons of the male
and the broad parafacial of both sexes.

Protocalliphora (P.) metallica (Townsend)
(Figs. 21, 39, 56, Map 11)

Protocalliphora splendida (Macquart).--Harbeck, 1907: 216 [N.
J., not rechecked; apparently first reference of splendida to
Protocalliphora; probably metallica but bicolor is possible].

Protocalliphora azurea (Fallén).--Gibson, 1917: 156 [Ottawa,
Ont.].Phormia metallica Townsend, 1919: 379[N.H., Va., Il.,
Wis., Wash.; females only in series; type locality, N.H.].

Phormia metallica.--Plath, 1919c: 373-8 [Wash.; ?possibly
cuprina, only females available].

Protocalliphora splendida Macquart.--Shannon and Dobroscky,
1924: 250 [Va. (part from type series of metallica); first
recognition of dichromatism].

P. splendida.--Johnson, 1925a: 216, in part [New England];
Dobroscky, 1925: 274-5 [N.Y.]; Shannon, 1926: 128 [In key];
Johannsen, 1928: 827 [N.Y.]; Johnson, 1932: 26 [Mass.]; Petch
and Maltais, 1932: 79 [Que.].

P. sp.--Thomas, 1936: 46, in part [Ohio].

P. splendida.--Judd, 1944: 146 [Que.]; Mason, 1944: 232-247
[Mass., mixture of metallica, sialia, shannoni].

186 Protocalliphora

Apaulina metallica.--Hall, 1948: 196-200 [Revision]; George and
Mitchell, 1948: 550, in part [N.Y].

Protocalliphora metallica.--Nolan, 1955: 55-60 [Ind.]; Stabler,
1959: 48 [Colo.]; Nolan, 1959: 355 [Ind.]; Boyes, 1961: 552ff.
[Cytology]; Boyes and van Brink, 1965: 543 [Cytology]; Hall,
1965: 926 [Catalog, widespread]; Stiner, 1969b: 1205-6 [Del.];
Boyes and Shewell, 1975: 446 [Mention of cytology].

P. sp. Ill.--Whitworth, 1976: 16ff.

Diagnosis. Dichromatic, the thorax and abdomen in male
shining metallic dark blue to bluish black, in female heavily gray
microtomentose and dull bluish to bluish green, with tergite 5 of
abdomen brightly cupreous to cupreous green; surstylus of male
long, curved and parallel-sided (Fig. 21).

Male. Parafacial and parafrontal silvery gray, often with a
very slight brownish tint. Body highly shining, dark blue to
bluish black, the thorax and abdomen concolorous, thorax darker
than abdomen and with three broad blackish blue stripes weakly
demarcated by the thin gray microtomentum, as viewed from
behind. Abdomen with thin gray microtomentum when viewed
from behind at a low angle. Calypteres opaque whitish, outer
rims at most tinted with yellow.

Frons narrow (Fig. 39), narrowing slightly anterior to ocellar
tubercle, at its narrowest 0.08 times head width (0.06-0.09) and
slightly broader than ocellar span (1.11x; 0.85-1.44):; parafrontal
strongly narrowed posteriorly, the prevertical area polished; one
row of parafrontal hairs that merge into the 2-3 irregular rows of
fine black parafacial hairs; parafacial obviously much wider than
width of 3rd antennal segment, | 1/2 times width of frons (1.51x;
1.33-2.00), 1.68 times ocellar span (1.43-2.00), and about 3/4
vibrissal interval (0.78x; 0.70-0.86); ocellar tubercle continued
forward as a sublinear acuminate polished area for a short
distance anterior to median ocellus, at least to level of uppermost
frontal bristles; cheek comparatively narrow, its height slightly
over 1/3 greatest length of eye (0.36x; 0.33-0.39).

Thorax without accessory notopleural bristles; tympanic
membrane and postalar wall usually bare. Fore tibia typically
with one posterior bristle, seldom with two and then usually only
on left or right, rarely on both.

Terminalia characterized by relatively long, moderately
narrow, curved but parallel-sided surstylus (Fig. 21); cerci
distally acuminate and parallel (as in Fig. 5b); aedeagus as usual
for the genus (as in Fig. 6a).

P. metallica 187

Female. Thorax and abdomen except Sth tergite blue to
bluish green, dull to weakly shining, heavily gray
microtomentose, 5th tergite bright shining cupreous or cupreous
green; scutum viewed from behind with three narrow shining
Stripes strongly demarcated by gray microtomentum; abdomen
with tergites 2 to 4 heavily gray microtomentose, 5th tergite very
thinly so and highly shining, thus adding to the contrast in color;
in extremely teneral specimens the 5th tergite shining greenish
with slightly coppery reflection; parafrontal and parafacial
brownish gray microtomentose, upper half of latter, viewed from
directly above, appears to have a subtriangular dark brown spot,
the spot changing to bright gray or yellowish gray when viewed
from below; preocellar area dull or only subshining, heavily
microtomentose.

Frons at vertex over 1/4 width of head (0.27x; 0.25-0.29) and
3.58 times ocellar span (3.00-4.17); frontal vitta with few and
inconspicuous hairs, usually a narrow polished area adjoining
anterior margin of median ocellus, occasionally more extensively
polished; parafrontal moderately wide throughout though
narrowing slightly above, with one row of fine black hairs,
merging below with 2-3 irregular rows on the parafacial;
parafacial width almost twice width of 3rd antennal segment,
0.45 times width of frons (0.40-0.53), 1.59 times ocellar span
(1.43-1.91), and 0.82 times vibrissal interval (0.72-0.92); cheek
comparatively narrow, its height 0.36 times maximum length of
eye (0.32-0.39).

Apparent normal length of adults, 8-9 mm. Specimens as
small as 5.5 mm have been seen.

Egg. Length 1000 w (956-1020)/14; breadth 310 yw
(275-380)/14; cuticle white in color, with small ridges.

First instar. Length up to three’ millimeters.
Cephalopharyngeal skeleton 215 w in length. Prothoracic fringe
absent. Posterior region: stigmatal plates 18 wu (16-20)/4 in
diameter; peritreme absent; mesostigmatal spines absent;
hyperstigmatal spines 3 w (1-5)/20; anal lobes small. Dorsal
cuticle with numerous spines 6 (2-10)/10. Ventral cuticle:
spines of anterior patch typically two-pronged, 2 wu (1-6)/20;
medial band present, spines 3 us (2-6)/20; posterior band present.

Second instar. Cephalopharyngeal skeleton 428 U
(400-456)/4 in length. Prothoracic fringe 8 wm (4-12)/50.
Posterior region: peritreme closed; stigmatal plates 77
(69-86)/9 in diameter; button distinct; mesostigmatal spines
present; both hyperstigmatal and hypostigmatal spines numerous,
as tubercles, 3 (2-5)/50. Dorsal cuticle with numerous spines,

188 Protocalliphora

14 pw (4-10)/50. Ventral cuticle: spines of anterior patch 7
(6-8)/50; medial band pronounced, spines 9 pw (2-10)/50;
posterior band present.

Third instar. Peritreme variable, usually weakly closed;
cuticular spines as tubercles, numerous between stigmatal plates
and in hypostigmatal area. Prothoracic fringe 37 wb (25-49)/100.
Cephalopharyngeal skeleton 730 uw (610-845)/78 in length.

Puparium. Length 7.3 mm (4.8-9.2)/90; breadth 3.3 mm
(2.0-4.2)/90; thin-walled, shiny, semi-translucent, brown.
Posterior region (Fig. 56a,b): stigmatal plates 160 4 (130-195)/192
in diameter; distance between buttons 560 uw (300-755)/96, and
across stigmatal plates 1185 u (690-1470)/96; stigmatal ratio 0.47
(0.41-0.48)/96; stigmatal area outlined in large plaques;
mesostigmatal spines, as tubercles, scarce, but present;
mesostigmatal folds absent; mesostigmatal plaques present;
hyperstigmatal spines as small tubercles, 4 wu (1-7)/100;
hypostigmatal area highly variable, ranging from folds absent
(Fig. 56a) to folds numerous and prominent (Fig. 56b);
hypostigmatal spines numerous, as tubercles, 3 uw (1-7)/100;
circumstigmatal folds highly variable, ranging from concentric
(Fig. 56a) to rectangular (Fig. 56b) and from pronounced to
virtually absent. Dorsal cuticle (Fig. 56c) with numerous minute
tubercles, 6 mw (4-14)/100; cuticular ridges weak (in eastern
forms), to virtually absent (in western forms). Ventral cuticle
(Fig. 56d): ventral band ratio 0.62(0.42-0.66)/100; spines of
anterior patch 7 w (2-12)/100; medial band pronounced, spines
6 pw (2-12)/100; posterior band pronounced.

Type Series. Holotype female, Franconia, N.H. (Mrs.
Slosson) [USNM], identified by Coquillett as P. splendida. This
is the only specimen of the type series that is marked in the
collection, although the species was based on a series of 28
females, "eight pinned from New Hampshire, Virginia, Illinois,
Wisconsin," and twenty (10 in alcohol) from Puget Sound, Wash.,
reared by O. E. Plath. Undoubtedly the old examples of
metallica from Milwaukee, Wis., Carlinville, Ill., and Rosslyn,
Va., were part of the original series, and they have been labeled
as probable paratypes. The three New Hampshire and three of
the four Puget Sound specimens cited by Townsend as a color
form were used by Shannon and Dobroscky (1924) as the type
series of aenea (N.H.) and hirudo cuprea (Wash.), respectively.
One Puget Sound specimen [USNM] and one labeled "ex nest
Western Robin" "Seattle, Wn., O. E. Plath"/Protocalliphora
splendida Macq." [det. Shannon] [Cornell U.] are metallica but
there is no indication whether these were part of the type series.

P. metallica 189

Specimens Examined. 469 males, 417 females.

Distribution. Widespread, British Columbia to Quebec and
Maine, south to California, Colorado, Iowa, and Virgina. April
19 (Conn.) and 21 (Iowa) to Oct. 10 (Ottawa, Ont.), dates of
hand-collected specimens.

Canada: Alberta (Banff), British Columbia (Abbotsford,
Kamloops, Oliver), Manitoba (Ninette near Brandon), Ontario
(Algonquin Park, Dundas, Low Bush (Abitibi), North Bay,
Osgoode, Ottawa, Toronto), Quebec (Armstrong, Aylmer, Kirk’s
Ferry, La Trappe, La Verendrye Provincial Park, Montreal,
Parkins Mills, Poltimore).

Contiguous United States: Alabama (Auburn, larvae and
puparia only), California (Devil’s Basin at 8200 ft, Mt. Shasta,
Yosemite Park, and Mariposa, Plumas, San Bernardino, and
Trinity Counties), Colorado (Colorado Springs), Connecticut
(Colebrook, Milford, New Haven, Stafford), Delaware (Newark),
Idaho (Moscow, Pocatello Valley in Oneida Co.), Illinois
(Carlinville, Downer’s Grove in DuPage Co., St. Joseph in
Champaign Co., Rockford), Indiana (Bloomington, Lafayette),
Iowa (Ames, Des Moines, Lodges State Park in Boone Co.),
Maine (Stetson in Penobscot Co., Southwest Harbor), Maryland
(College Park, Cumberland), Massachusetts (Bristol Co.,
Brookline, Forest Hills, Groton, Horse Neck Beach, Martha’s
Vineyard, North Andover, Swampscott), Michigan (Bath, East
Lansing, Midland, Owosso, and Pentwater, and Bay, Hillsdale,
and Livingston Counties), Minnesota (Ft. Snelling, Grand Rapids,
and Hennepin, Ramsey, and St. Louis Counties), Nebraska
(Omaha), New Hampshire (Coos Co., Durham, Franconia,
Peterboro), New Jersey (Clementon, Mountain Lakes, West
Orange), New York (Ithaca, and Erie, Essex, Franklin, Genesee,
and Tompkins Counties, also Babylon, Roslyn, and Mastic on
Long Island and Watchogue on Staten Island), Ohio (Columbus,
Lakewood, Sandusky, and Wadsworth, and Franklin and Union
Counties), Oregon (Coos Bay, Marshfield, Mt. Hood),
Pennsylvania (Butler, Pittsburgh, State College, Wind Gap),
South Dakota (no locality), Utah (Benson in Cache Co.,
Brighton, Box Elder Co.), Vermont (Manchester), Washington
(Asotin Co., Mt. Rainier, Puget Sound, Spokane), West Virginia
(Cranberry Glades), and Wisconsin (Milwaukee).

Basis of description of immature stages: Egg (14 specimens):
Algonquin Park, Ont., from caged female, July 1955. Larvae, Ist
instar (2 specimens): Algonquin Park, Ont., red-winged
blackbird, June, 1954. Second instar (5 specimens): same data.
Third instar (20 specimens): Algonquin Park, Ont., red-winged

190 Protocalliphora

blackbird, June, 1955 (7); yellow warbler, June, 1955 (3); song
sparrow, July, 1955 (3); slate-colored junco, July, 1954 (7).
Puparium (96 specimens): Algonquin Park, Ont., red-winged
blackbird, June, 1955 (27); common grackle, June, 1955 (26);
yellow warbler, July, 1955 (10); song sparrow, August, 1955 (10);
gray catbird, July, 1955 (13); swamp sparrow, July, 1953 (10).

Hosts (Eastern). Red-winged blackbird, indigo bunting,
northern cardinal, gray catbird, yellow-breasted chat, alder
flycatcher, common grackle, slate-colored junco, American
robin, a number of sparrows (field, grasshopper, savannah, song,
swamp, vesper, and white-throated), American tree swallow,
brown thrasher, rufous-sided towhee, prairie and yellow
warblers; also, from puparia alone, black-billed cuckoo,
American goldfinch, eastern kingbird, chipping sparrow, bank
and northern rough-winged swallows, scarlet tanager, Nashville
warbler, northern waterthrush, a wren (Alabama), and common
yellowthroat.

Hosts (Western). Brewer’s and red-winged blackbirds, lazuli
bunting, blue grosbeak, fox sparrow, and an unidentified sparrow
(Utah).

For the scarlet tanager record, no specimens were seen but
the words of Dobroscky (1925) assure us that the species was
metallica. She described the spines around the stigmatal area of
puparia as "so minute that they seemed like mere stipples."
Incidentally this is the only record known to us of
Protocalliphora from that host.

The statement by Van Tyne and Berger (1976) that "54
species of birds are known to be parasitized by P. metallica is
erroneous, apparently stemming from confusion with the list of
54 species and subspecies cited by Hall (1948) as known hosts at
that time of all species of Protocalliphora in North America. At
present we have records of 35 species of hosts of metallica.

Ecology and Biology. P. metallica primarily infests nests
close to the ground or at shrub level, particularly in open fields
or marshes, and including cavity nests close to the ground. The
biology of this common species has been investigated in some
detail by Bennett at Algonquin Park, Ont. (Bennett 1957).

This is the species most often collected by hand, whereas
most species have been taken for the most part by rearing from
larvae or puparia found in birds’ nests. It is also most often
collected at flowers, especially at goldenrod in the autumn and
fall. Both of these points may rest on the facts that it is an
extremely common species and frequents low vegetation levels
where it would more often be noticed.

P. metallica 191

Variation. A few specimens showed one to four fine hairs in
~ the tympanic pit (three on each side in the holotype itself), and
a few short black hairs on the postalar wall.

Four males, one female from Groton, Mass., provide a
warning note on color variation. Johnson (1932), who called
these splendida, noted that in the males "the terminal segment of
the abdomen is a bright green, not blue." When the original
series was found in the C. W. Johnson Collection now at Boston
University it proved to be metallica. The characteristic
"sranular-type" puparia were associated with the specimens. The
"green" color is more of a greenish blue, exaggerated by the more
shining appearance of the Sth tergite compared with the
gray-microtomentose preceding segments.

In small or immature females of metallica, the distal segment
of the abdomen may be shining green with only a faint coppery
tint. This variation is clearly shown in ten females reared from
a northern cardinal nest at East Falls Church, Va., in which only
the most mature specimens show the characteristically
copper-colored Sth tergite, sharply contrasted with the
preceding segments.

Variation in the parafacial hairs will cause some difficulty in
identifying isolated males. Small specimens seem sparsely haired
with relatively few hairs in two irregular rows, or occasionally a
partial third row, whereas large specimens have more hairs in
three or even almost four irregular rows. Inasmuch as metallica
is especially a parasite of small birds and many specimens are
undersized, this characteristic of relatively few hairs will serve
for most specimens.

As a possibly interesting check on variation in ratios, 25
males reared by Bennett from six different nests at Algonquin
Park, Ont., were measured and ratios calculated for comparison
with the 25 males on which were based the ratios used in
description. These 25 were a sample from the wide range of the
species, coming from 15 localities in 9 states and Ontario,
including only five specimens from five nests in Algonquin Park.
The following comparison reveals only slight differences:

192 Protocalliphora

General series Algonquin Park
frons: head 0.08 (.06-.09) 0.08 (.07-.10)

frons: ocellar span 1.11 (.85-1.44) 1.13 (1.00-1.36)
parafacial: frons 1.51 (1.33-2.00) 1.45 (1.17-1.83)
parafacial: ocellar span 1.68 (1.43-2.00) 1.65 (1.43-1.92)
parafacial: vibrissal interval 0.76 (.70-.86) 0.76 (.625-.88)
cheek: eye 0.36 (.33-.39) 0.35 (.32-.375)

Remarks. In older literature, metallica was synonymized
with splendida Macquart, but we concur with Hall (1948) in
regarding the latter as a species dubia. For a discussion of this,
see the heading Calliphora splendida at the end of the species
descriptions.

Records of or specimens identified as "splendida" cannot be
assumed to be metallica--although they often are--because there
is another northeastern species (bicolor) in which females have
tergite 5 coppery tinted, and because males identified as
splendida by older authors often prove to be sialia.

Identifications of metallica can also be suspect. The possible
confusion with bicolor has just been noted. In Plath (1919c),
"metallica" proved in part to be braueri, and in Neff (1945)
"metallica" proved to be cuprina. "P. metallica" of James (1955),
based on a male from Yosemite National Park, Calif., is
spatulata.

Females of metallica have long been regarded as the most
distinct in the genus because of the shining cupreous 5th tergite
contrasting with the blue or bluish purple of the 2nd to fourth
tergites. Unfortunately this can now lead to misidentifications
because two other species, bicolor in the East and cuprina in the
West, have the same feature and the females of the three species
are very similar and will be difficult to distinguish. The key to
females utilizes a minor feature of the microtomentum of the
scutum which hopefully will separate females of metallica from
the others.

Males of metallica have no outstanding or peculiar
characteristic or combination of characteristics. The surstylus is
curved, placing the species in that group (See outline of groups,
preceding the keys, and the key to males).

Puparia of metallica are distinctive because of the "fine
sandpaper" appearance imparted by the short tuberculose spines.
Puparia of metallica are similar to those of deceptor, interrupta,
and shannoni, but separated from shannoni by the tubercle-like
spines of the hyperstigmatal area and from deceptor by the
shorter tubercle-like spines of the dorsal and ventral cuticle.

P. parorum = 193

Although similar in many respects to interrupta, the latter species
may be separated by the interruption of the medial ventral band
and the markedly smaller ventral band ratio. Puparia of
metallica may be separated from braueri by the presence of the
three distinct ventral spine bands.

Protocalliphora (P.) parorum, new species
(Figs. 22, 57, Map 10)

Protocalliphora splendida var. near hirundo.--Storer, 1927: 227
[From nest of mountain chickadee, San Bernardino Mts.,
Calif.; material lost, possibly parorum, considering the host].

P. sp. VI.--Whitworth, 1976: 16ff. [Utah].

P.n.sp. no. 1.--Gold and Dahlsten, 1984: 560-572 [Calif.; effect
on nestlings of chestnut-backed and mountain chickadees].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic bluish to bluish purple; parafrontal and parafacial dark
gray microtomentose; calypteres white; surstylus narrow and
curved.

Male. Head darker than usual in Protocalliphora, black in
ground color with parafrontal and parafacial dark gray
microtomentose. Body shining metallic bluish to bluish purple,
scutum partly gray microtomentose, especially anterior to
transverse suture, with two broad stripes of microtomentum
demarcating three broad shining stripes, as viewed from behind.
Abdomen shining in direct view, tergites 2-4 thinly gray
microtomentose when viewed from behind at a low angle.
Calypteres white.

Frons relatively narrow, at narrowest, anterior to median
ocellus, 0.08 times width of head (0.06-0.10) and 1.13 times
ocellar span (0.91-1.42); ocellar tubercle with none to few bristles
or bristlelike hairs except for ocellar bristles; preocellar area
narrowly triangular to acuminate, dark gray microtomentose,
slightly shining, sometimes rugose; frontal bristles and
parafrontal hairs usually ending well in advance of level of
median ocellus, uppermost frontals weak and hairlike; parafrontal
with one to two irregular rows of hairs on lower half, strongly
narrowed dorsally, its microtomentum narrowed to pinched out,
and prevertical area usually polished; parafacial with numerous
coarse black hairs in 3-4 irregular rows, no changeable spot along
upper margin; parafacial 1.41 times width of frons (1.18-1.70),
1.60 times ocellar span (1.43-1.83), and 0.77 times vibrissal

194 Protocalliphora

interval (0.69-0.86/21); height of cheek 0.42 times maximum
length of eye (0.38-0.45); occiput with about three irregular rows
of black hairs behind postocular setae.

Thorax lacking accessory notopleural bristles; tympanic
membrane and postalar wall with none to few short, pale and
inconspicuous hairs. Fore tibia typically with one posterior
bristle.

Terminalia: cerci long and narrow to acute apices (cf. Fig.
5b); surstylus narrow and curved (Fig. 22); aedeagus as usual in
the genus (cf. Fig. 6a).

Female. Concolorous with male, but scutum much more gray
microtomentose, especially posterior to transverse suture; head
dark gray microtomentose to slightly brownish tinted. Frons 0.26
times head width (0.25-0.28) and 3.33 times ocellar span
(2.875-3.71); preocellar area dull, finely and _ evenly
microtomentose; parafrontal above tending to be _ thinly
microtomentose, subshining; parafacial as in male, densely haired
without changeable spot, width 0.47 times that of frons
(0.43-0.52), 1.55 times ocellar span (1.25-1.71), and 0.81 times
vibrissal interval (0.69-0.92/24); height of cheek 0.43 times
maximum length of eye (0.40-0.47).

Apparent normal length of adults, 8-9.5 mm.

Third instar. One specimen examined, additional
measurements derived from puparia. Prothoracic fringe 80 pu
(50-120)/70. Cephalopharyngeal skeleton 950 w (800-1050)/15
in length.

Puparium. Length 7.4 mm (7.2-7.5)/5; breadth 2.7 mm
(2.4-3.3)/10; thick-walled, dull, brown. Posterior region (Fig.
57a,b): stigmatal plates 160 wu (150-175)/20 in diameter; distance
between buttons 565 uw (500-625)/10, and across the stigmatal
plates 1100 uu (1000-1200)/10; stigmatal ratio 0.52 (0.45-0.58)/10;
stigmatal area outlined in folds and a few plaques; mesostigmatal
folds moderately pronounced (Fig. 57a) to virtually absent (Fig.
57b), the former bisecting the stigmatal region; hyperstigmatal
spines numerous, 33 (8-48)/100; hypostigmatal area with
numerous, pronounced, irregular folds; meso- and hypostigmatal
spines sparse, the latter 15 us (8-30)/100; circumstigmatal folds
absent (Fig. 57b) to pronounced concentric (Fig. 57a). Dorsal
cuticle (Fig. 57c) with numerous, large spines, 35 w (15-55)/100;
cuticular ridges weak. Ventral cuticle (Fig. 57d): ventral band
ratio 0.74 (0.58-0.87)/20; anterior band pronounced, anterior
patch distinct, spines 15 us (8-30)/100; medial band pronounced,
spines 24 wu (15-50)/100; posterior band pronounced.

P. parorum =-195

Type Series. Holotype male, allotype, and 16 paratypes (1
male, 15 females), California: Modoc Co., 7 mi N Adin, June 19,
1968, ex chickadee nest (D. L. Dahlsten). Holotype and allotype
in the California Academy of Sciences, paratypes in the
California Academy of Sciences, U.S. National Museum of
Natural History, and University of California at Albany (Division
of Biological Control). Additional paratypes: California [CAS,
USNM, and U. Calif., Albany]: 5 males, 5 females, Contra Costa
Co., Wildcat Canyon, June 1981, ex nest chestnut-backed
chickadee (C. Gold); male, 3 females, El Dorado Co., U.
California Blodgett Forest, 8 mi E Georgetown, nest of brown
creeper, emerged July 6 and 11, 1978 (C. Gold); 6 males, 7
females, Blodgett Forest, emerged from nest of chestnut-backed
chickadee May 29-July 11, 1978 (C. Gold); 2 males, 12 females,
Blodgett Forest, ex nestboxes of mountain chickadee, July 1-22,
1978 (C. Gold); male, 20 females, Blodgett Forest, July 30-Aug.
6, 1977, ex nestboxes of chestnut-backed chickadee (C. Gold); 4
males, 18 females, Blodgett Forest, July 25-Aug. 6, 1977, ex
nestboxes mountain chickadee (C. Gold); 3 females, Blodgett
Forest, July 21, 1979, ex nestbox chestnut-backed chickadee (C.
Gold); male, Lassen Co., Fredonyer Peak, 25 mi N Susanville,
July 25, 1967, ex chickadee nest (F. M. Stephen); 8 males, 13
females, same locality as preceding, July 4, 1969, ex nest of
red-breasted nuthatch (D. L. Dahlsten); female, Los Angeles Co.,
Tanbark Flat, June 22, 1950 (H. F. Robinson) [U. Calif., Davis];
24 males, 49 females, Modoc Co., various localities (Roney Flat,
Tom’s Creek, Yellowjacket) and dates in 1978, chiefly July, ex
nests of mountain chickadee (C. Gold); 22 males, 30 females,
Modoc Co. near Adin, emerged July 25-Aug. 8, 1977, ex
nestboxes mountain chickadee (C. Gold); 10 males, 9 females,
Mono Co., Pilot Springs, 23 mi E Lee Vining, July 16, 1968, ex
nest mountain chickadee (D. L. Dahlsten); 10 males, 12 females,
Mono Co., 27 mi E Lee Vining, July 17, 1968, ex nest mountain
chickadee (D. L. Dahlsten). Utah: 4 males, 1 female, Smithfield,
July 11, 1970, nest of black-capped chickadee (T. L. Whitworth)
[USNM, Whitworth Colln.].

Non-paratype Material: One female, Utah, and one male,
Utah, Box Elder Co., July 21, 1969 (T. L. Whitworth, Malaise
trap, and 21 males, 32 females, various localities in California,
duplicated in the type series. Also 3 males, 6 females, California:
Inyo National Forest, Sentinel Meadows, Aug. 6, 1963 (S. G.
Herman) [U. Calif., Berkeley]. Also two isolated specimens, a
male, California: Modoc Co., Adin Bridge, emerged July 27,
1978, ex barn swallow nest, and a female, probably parorum,

196 Protocalliphora

California: Lassen Co., Summit Camp, June 28, 1949 (Claude I.
Smith) [U. Calif., Berkeley].

Specimens Examined. 126 males, 239 females.

Distribution. California and Utah.

Basis of description of immature stages: Larva, 3rd instar (4
specimens): Modoc Co., Calif., mountain chickadee, coll. Gold.
Puparium (10 specimens): Modoc Co., Calif., mountain chickadee.

Hosts. Black-capped, chestnut-backed, and mountain
chickadees (chiefly), brown creeper (few), red-breasted nuthatch
(few), and barn swallow (one). Whitworth (1976) also recorded
it from Oregon junco and house wren in Utah, and Gold and
Dahlsten (1981, 1984) recorded it from Bewick’s wren.

Ecology and Biology. The extensive available material was
reared almost exclusively from nests of chestnut-backed and
mountain chickadees, but finding specimens in Utah suggests a
wider range when chickadee nests are investigated in other parts
of the West. Gold and Dahlsten (1984) reported on the effects of
parorum on chickadee nestlings. Cavity nesters in forest habitats
appear to be the principal hosts.

Variation. Of the 360 specimens available, 347 have only one
posterior bristle on the fore tibia. None have two on each fore
tibia. Five males, four females show two bristles on one side or
the other, but even most of these are not the usual well spaced
bristles of species that characteristically have two. In some
specimens the upper bristle is very short compared with the lower
bristle, which is the standard one, and in two indivduals the two
bristles are close together near the usual position of the lower
bristle, not well spaced. Clearly, one posterior bristle is the
regular character state for parorum.

Remarks. P. parorum is one of the complex of species with
males characterized by a narrow curved surstylus. Single
specimens will usually be difficult or impossible to identify with
certainty. In the present case, the species is especially dark, a
dark gray microtomentum in both sexes that gives it a different
habitus from the other species, although this is not easily
expressed in a key for those who do not have authoritatively
identified examples before them. The three most difficult
species to distinguish are parorum, fallisi, and tundrae, and the
reader is referred to the general key. The first and last each have
a characteristic habitus, once one is accustomed to the usual
appearance of these flies. In addition, P. tundrae occurs only in
the far north, in northern Canada and Greenland, as far as
known at present, and parorum seems typically a parasite of
chickadees, although some other small birds are known also as its

P. sapphira 197

hosts. Moreover the apparent frequency of chickadees as hosts
of parorum could have been enhanced as a result of concentrated
study on the chickadees by Dahlsten and colleagues.

Puparia of parorum can be confused with those of bicolor and
cuprina. They can be separated from bicolor by the larger ventral
band ratio and from cuprina by the larger prothoracic fringe.
Great care should be taken with these species and a general
evaluation of all available characters made before assigning a
specific designation.

Storer’s (1929) record of Protocalliphora splendida var. near
hirundo from the nest of a mountain chickadee in the San
Bernardino Mts., California, probably referred to parorum. The
material is apparently lost (personal communication from
Professor Storer). Storer recorded 76 larvae and 24 puparia in the
nest, at 7400 ft altitude.

Etymology. The specific name is in the genitive plural, from
the predominance of parasitism on species of the genus Parus, the
chickadees.

Protocalliphora (P.) sapphira (Hall)
(Fig. 26)

Protocalliphora splendida var. sialia.--Shannon and Dobroscky,
1924: 252, in part [Naknek Lake, Alaska].

Apaulina sapphira Hall, 1948: 200 [Naknek Lake, Alaska; type
series included in sialia in Shannon and Dobroscky 1924].

Protocalliphora sapphira.--Hall, 1965: 926 [Catalog].

Diagnosis: Sexes concolorous, monochromatic, shining
metallic dark bluish; calypteres white; front tibia with 2 posterior
bristles; frons of male relatively broad, 0.17 times head width;
surstylus narrow and strongly curved.

Male (only holotype available). Body shining, dark metallic
bluish; thorax and abdomen concolorous, former slightly darker,
and duller with gray microtomentum between three broad shining
Stripes as viewed from behind, the gray stripes equal to or wider
than the shining stripes; abdomen thinly microtomentose but
highly shining, with linear black to purple median stripe that is
especially evident on tergites 3 and 4 as viewed from behind at
a low angle. Calypteres white, rims pale yellow.

Frons relatively broad for the genus, not narrowing anterior
to median ocellus, at vertex 0.17 times head width and 1.9 times
ocellar span; parafrontal and parafacial with light-brown-tinted

198 Protocalliphora

microtomentum and sparse black hairs, width of parafacial 1.8
times breadth of 3rd antennal segment, 0.85 times width of frons,
1.6 times ocellar span, and 0.79 times vibrissal interval (not equal
as stated in Hall’s description; the ptilinal fissure is not
completely closed in the holotype and the true lunule width is
narrower); occipital hairs predominantly black above, in
approximately three irregular rows; cheek relatively broad, its
height 0.46 times maximum length of eye.

Thorax without accessory notopleural bristles; postalar wall
not conspicuously haired, with one to three or four short, weak
hairs visible under high magnification; tympanic membrane
weakly setose (four on left, three on right). Fore tibia with two
posterior bristles (a third bristle on right side is probably an
aberration).

Terminalia: cerci distally acuminate, parallel (as in Fig. 5b);
surstylus narrow, parallel sided, strongly curved, distally truncate
(Fig. 26); aedeagus as usual for genus (cf. Fig. 6a).

Female (two paratypes). Color as in male; microtomentum of
frons and parafacial more obviously brownish or brownish
yellow; parafacial brownish above but silvery below. Frons at
vertex 0.30-0.31 times head width and 1.73 times ocellar span,
strongly broadening anteriorly, entirely microtomentose in
preocellar area; reclinate orbital bristles present; parafacial broad,
nearly twice breadth of 3rd antennal segment, over half (0.54)
width of frons, 1.73 times ocellar span, and 1.08 times vibrissal
interval; cheek height 0.49 and 0.50 times maximum length of
eye. Fore tibia with two posterior bristles in one paratype, one
in other.

Length of adults, 7.5 mm. (This does not necessarily indicate
a small species; the few individuals might have been underfed).

Immature stages. Unknown.

Type Series. Holotype male and paratype female, Savonoski,
Naknek Lake (now Lake Coville, near base of Alaska Peninsula),
Alaska, Aug. 1, 1919 (J. S. Hine); paratype female, same locality
and collector, June 1918. These specimens were part of the
syntype series of P. sialia S. & D. (1924:252). Both Shannon and
Dobroscky (1924) and Hall (1948) stated that all three specimens
were collected Aug. 1, 1919; the different date on one of the
paratypes was apparently overlooked. Type series in the U.S.
National Museum of Natural History. There is also a female,
same locality, Aug. 13, 1919 (A. J. Basinger) [CAS], which was
not included in the type series.

Specimens Examined. | male, 3 females.

Distribution. Alaska, as far as known.

P. sapphira 199

Hosts, and Ecology and Biology. Unknown.

Variation. The extreme variability in the short type series
makes it impossible to properly assess the typical characteristics
of the species. One paratype female has 2 posterior bristles on
the fore tibia and a fairly good tympanic tuft of 6-8 hairs on
each side, but only 2 or 3 very weak hairs on the postalar wall.
The other paratype female has only one posterior bristle on the
fore tibia, a fairly good tympanic tuft of 10-12 conspicuous
black setae, but few (4) weak and inconspicuous hairs on the
postalar wall. The holotype has two posterior bristles on the fore
tibia, but both tympanic membrane and postalar wall only weakly
haired (see description).

Remarks. In the holotype, the ptilinal fissure had not
completely closed and the anterior part of the frons is not
completely sclerotized. The specimen otherwise does not appear
teneral and the upper half of the frons appears normal, hence the
measure of the frontal width is accepted. This wide frons
distinguishes sapphira from males of all other species having
curved and narrow surstylus.

Females are not distinctive. The number of posterior bristles
on the fore tibia probably varies, and further material is needed
to show whether it is typically a two-bristle species, or whether
the reverse is true. Our experience indicates that either is
possible in these northern species, but that two bristles is the
most common characteristic. The adequate series in tundrae
demonstrates that even in a species where the two-bristle state is
common, there will be a sizeable proportion of specimens that
have only one bristle on the fore tibia.

The setose tympanic membrane and two posterior bristles on
the fore tibia relate sapphira to Palearctic species, as might be
expected from its far northern range, but it has male terminalia
distinct from any Palearctic species. The chaetotactic characters
just noted apply also to chrysorrhoea, in which the hair tufts are
larger and stronger, but the wide frons and narrow and curved
surstylus easily separate sapphira. The surstylus in profile is
suggestive of metallica, but the latter is a much different species
that would not be confused. In some respects, and especially in
females, sapphira resembles spatulata, but the latter has,
regularly, one posterior bristle on the fore tibia and no tympanic
setae, and in the male conspicuously polished prevertical area.

Hall stated that sapphira differed from all other Apaulina in
having apical bristles on the scutellum. In reality sapphira as
well as most Protocalliphora lack well developed apical bristles
although one pair of hairs is slightly stronger than surrounding

200 Protocalliphora

hairs and sometimes, especially in large specimens, does appear
bristlelike. In most specimens, this apical pair is actually
subapical in varying degrees, less so in sapphira than in most
species.

The only published record of P. sapphira other than those
listed in the synonymy is that by Sailer and Lienk (1951), but
later study showed these specimens to be P. chrysorrhoea
(Meigen). The "cliff swallow" host was an error for bank swallow
(R. I. Sailer, personal communication).

Protocalliphora (P.) seminuda, new species
(Figs. 25, 58, Map 6)

Protocalliphora chrysorrhaea [sic] (Meigen).--Walton, 1914: 175,
in part [Koehler, N. Mex., mixed with P. braueri).

P.n. sp. A.--Horning and Barr, 1970: 73 [Idaho].

Protocalliphora sp. X.--Whitworth, 1976: 16ff. [Utah].

Diagnosis. Sexes concolorous, monochromatic, shining dark
blue to bluish purple; calypteres whitish; male frons moderately
narrow; parafacial and parafrontal of male conspicuously silvery
microtomentose, with few hairs; surstylus in profile curved
though not slender.

Male. Head with microtomentum of parafrontal and
parafacial silvery. Body dark bluish to bluish purple, the thorax
and abdomen concolorous, shining, the abdomen highly so, with
thin gray microtomentum visible only from behind at an
extremely low angle, the dorsum of thorax duller, more obviously
microtomentose, especially on presutural area of scutum, the
scutum viewed from behind with three broad shining stripes.
Calypteres whitish, outer rims slightly pale-yellow tinted.

Frons moderately narrow, narrowing only slightly anterior to
ocellar triangle, at narrowest 0.08 times (0.06-0.09/9) head width
and barely wider than ocellar span (1.16x; 1.00-1.50/9);
preocellar area shining, narrowly acuminate; frontal vitta at its
narrowest less than twice width of a parafrontal; frontal bristles
strong below, rapidly decreasing in strength above, the uppermost
short, fine and hairlike; parafrontal silvery microtomentose
dorsad nearly to median ocellus, and almost bare of hairs except
for four or five in one row near lower frontal bristles (in one
specimen a complete but inconspicuous row close to frontal row),
prevertical area shining black; parafacial moderately wide, its
width greater than breadth of 3rd antennal segment, 1.50 times

P. seminuda 201

(1.33-1.83/9) width of frons, 1.74 times ocellar span
(1.36-2.08/9), and 0.90 times (0.75-0.92/7) vibrissal interval,
thinly haired along anterior half in about two to three irregular
rows, posterior half bare; height of cheek 0.38 times
(0.36-0.41/9) maximum length of eye.

Thorax without accessory notopleural bristles; tympanic
membrane and sometimes postalar wall with a few scattered
hairs. Fore tibia with one median posterior bristle.

Terminalia resembling those of metallica, the cerci straight
and parallel, deeply notched, long, slender and acute at apex (cf.
Fig. 5b); surstylus in profile curved, though not slender (Fig. 25);
aedeagus as usual for the genus (cf. Fig. 6a).

Female. Microtomentum of parafacial and parafrontal light
brownish yellow tinged. Body color as in male but mesonotum
somewhat duller, with more abundant grayish microtomentum.
Frons broad, about 1/3 width of head (0.33x; 0.32-0.36/6) and
four times ocellar span (3.56-4.43/6); frontal vitta on each side
often with several coarse setulae mesad of the frontal bristles
(broken off in allotype and one paratype, but sockets evident);
preocellar area broad, dull, subshining but evenly brownish gray
microtomentose; parafrontal sparsely haired, in about two
irregular rows, microtomentose throughout but prevertical area
thinly so and subshining; parafacial fairly broad, 0.41 times
(0.39-0.45/6) width of frons, 1.67 times ocellar span
(1.50-1.875/6), and 0.92 times vibrissal interval (0.88-1.00/3),
sparsely haired on mesal half, bare on outer half; height of cheek
0.44 times (0.42-0.48/6) maximum length of eye.

Apparent normal length of adults, 8-9.5 mm.

Third instar. No specimens examined, characters derived
from puparium. Prothoracic fringe 50 pm (40-65)/26.
Cephalopharyngeal skeleton 860 (800-950)/6 in length.

Puparium. Length 7.9 mm (7.4-8.3)/6; breadth 3.4 mm
(3.1-3.7)/7; medium thick-walled, dull, brown. Posterior region
(Fig. 58a): stigmatal plates 170 ww (150-175)/14 in diameter;
distance between buttons 590 wu (575-625)/7, and across stigmatal
plates 1175 w (1000-1375)/7; stigmatal ratio 0.51 (0.40-0.63)/7;
stigmatal area outlined in plaques and folds; mesostigmatal area
with numerous pronounced folds and plaques, bisecting the
stigmatal region; hyperstigmatal spines numerous, 22 u (8-40)/70;
hypostigmatal area rugose with numerous, pronounced, irregular
folds; hypostigmatal spines scarce, 14 wm (18-25)/70;
circumstigmatal folds pronounced; concentric. Dorsal cuticle
(Fig. 58b) with numerous spines, 22 mu (15-40)/70; cuticular
ridges absent or if present, very weak. Ventral cuticle (Fig. 58c):

202 Protocalliphora

ventral band ratio 0.63 (0.51-0.67)/9; anterior band pronounced,
anterior patch distinct, with numerous small spines, 12 wu
(8-25)/70; medial band pronounced, spines 20 pw (15-30)/70;
posterior band pronounced on all segments.

Type Series. Holotype male, and allotype, Koehler, N. Mex.
(W. R. Walton; Webster No. 7707; reared from horned lark).
Type in the National Museum of Natural History [USNM].
Paratypes: 3 males, 2 females, same data as holotype [USNM]; 2
males, California: Goodale Creek, N of Lone Pine, Inyo Co.,
Apr. 3, 1953 (H. B. Leech) [CAS]; 2 males, Craters of the Moon
National Monument, Idaho, June 29, July 23, 1965 (D. S.
Horning, Jr.), both collected in a Malaise trap [CAS]; 3 males, 6
females, Pocatello Valley, Oneida Co., Idaho, July 5, 1970 (T. L.
Whitworth, nest of Brewer’s blackbird [Whitworth Colln.,
USNM]; 3 males, Benson, Cache Co., Utah, July 9, 1970 (T. L.
Whitworth), nest of barn swallow, in mixed infestation with P.
halli [USU]; male, Green Lake, Uinta Mts., Utah, Aug. 17, 1940
(D. G. Hall) [USNM]; male, female, Stratton Experimental
Watershed near Saratoga, Wyo., May 26-June 2, 1976 (J. M.
Schmid), from pit and Malaise traps in sagebrush [USNM].

Non-paratype Material: Male, Box Elder Co., Utah, July 9,
1969 (K. J. Capelle) [Whitworth Colln.]; male, Summit Co., Bear
River, Utah, Aug. 5-12, 1971 (W. Hanson, G. F. Knowlton,
Malaise trap) [Whitworth Colln.].

The holotype and the Craters of the Moon paratypes are of
the usual size for most Protocalliphora, and we assume that this
is the normal size for the species. The other specimens are in
general smaller, and some are more or less teneral, especially the
New Mexico specimens, most of which have the abdomen more
or less collapsed.

Specimens Examined. 18 males, 10 females.

Distribution. California, Idaho, New Mexico, Utah,
Wyoming.

Basis of description of immature stages: Puparium (7
specimens): Koehler, N. Mex., horned lark (3); Pocatello Valley,
Oneida Co., Idaho, Brewer’s blackbird (4).

Hosts. Brewer’s blackbird, horned lark, and barn swallow.

Ecology and Biology. Two few hosts are known to give any
reliable picture of the host associations. Walton (1914), as already
discussed under braueri, stated that the larvae of P.
"chrysorrhaea" [sic], as it was then identified, "were contained in
purulent sores on the sides of the body near the legs and on the
neck." The embedding of the larvae in the skin is well known for
braueri and perhaps it is also true for seminuda. Walton’s brief

P. shannoni =. 203

published notes say nothing of rearing any larvae from the nest,
but rather they suggest that all maggots were taken directly from
a weak and emaciated fledgling. Possibly some larvae or puparia
may have been found in the nest material and reared as if all
were the one series. Certainly the infestation was mixed; braueri
and seminuda are decidedly different species and cannot be
confused in either sex, nor in the mature larvae and puparia.

Variation. The allotype has the chaetotactic aberration of
four orbital bristles on the left side and three on the right.

Remarks. The head of the male is similar in proportions and
silvery microtomentum to that of sialia--indeed the New Mexico
series stood in the collection under sialia and was so identified
by Shannon and Dobroscky (1924)--but the parafacials are
narrower and have distinctly fewer hairs, and the parafrontals are
almost bare. The male terminalia resemble those of metallica and
fallisi. For males, the thinly haired and partly bare parafacials
are distinctive, also the narrow frons with very narrow frontal
vitta. The females are not well separated from those of hirundo
and other western species. The coarse setulae towards the outer
margins of the frontal vitta might offer a means of recognition,
but the sample is too small to be sure of the consistency of the
character.

Puparia of seminuda can be confused with those of hesperia,
hesperioides, and spenceri, which they broadly resemble. They
can be distinguished, however, from hesperioides on the basis of
the shorter prothoracic fringe, and from spenceri on the basis of
the concentric circumstigmatal folds and absence of dorsal
cuticular ridges.

Etymology. The specific name, an adjective, is compounded
from the Latin prefix semi, half or partly, plus nudus, bare.

Protocalliphora (P.) shannoni, new species
(Figs. 13, 59, 60, Map 12)

Protocalliphora sialia, in part.--Shannon and Dobroscky, 1924:
252 [White Mts., N.H., Farmingdale and Ithaca, N.Y.].

P. splendida form sialis [sic].--Johnson, 1925a: 216 [Peterboro,
N. H. specimen].

P. splendida splendida.--Johnson, 1929a: 29 [Ashland, N.H.,
from robin nest].

P. splendida.--Mason, 1944: 232-247, in part [Mass.].

204 =‘ Protocalliphora

P. sp. (or n. sp.) near sialia.--Judd, 1954: 123 and 1957: 32
[London, Ont.]; Boyes, 1961: 557 [Cytology]; Boyes and van
Brink, 1965: 543 [Cytology]; Boyes and Shewell, 1975: 442,
446 [Cytology].

P. sp. VII.--Whitworth, 1976: 16 ff. [Utah].

Diagnosis. Sexes concolorous, monochromatic, shining dark
metallic blue to bluish black; calypteres white; frons of male very
narrow; surstylus digitate, of intermediate width.

Male. Head black in ground color; parafrontal and parafacial
gray microtomentose, bright but not silvery, rather a slightly pale
brown tint. Thorax shining dark metallic blue to bluish purple
or bluish black, dorsum thinly microtomentose, especially
anteriorly, the microtomentum not demarcating stripes and
scutum appearing entirely shining to the unaided eye. Abdomen
concolorous with thorax, but generally a brighter blue, sometimes
tinged with green, and very shining, thinly microtomentose when
viewed from behind at a low angle. Calypteres opaque whitish,
at most the outer rims tinged with yellow.

Frons very narrow, 0.06 times width of head (0.05-0.07), and
equal to (or nearly so) ocellar span (0.985x; 0.83-1.18));
parafrontal with only one row of hairs, on lower fourth, merging
with parafacial hairs of which there are two or sometimes three
irregular rows; parafacial comparatively narrow, equal to or
barely wider than 3rd antennal segment, nearly twice width of
frons (1.78x; 1.46-2.20), 1.76 times ocellar span (1.50-2.20), and
0.70 times vibrissal interval (0.58-0.815); cheek relatively narrow,
about 1/3 maximum length of eye (0.37x; 0.34-0.40).

Thorax without accessory notopleural bristles and without
hairs on tympanic membrane, and with only an occasional hair or
two on postalar wall. Fore tibia with one posterior bristle.

Terminalia: cerci distally acuminate, narrow and parallel;
surstylus digitate, of intermediate width (Fig. 13), somewhat as
in sialia, broadly rounded at distal end; aedeagus as usual in the
genus (cf. Fig. 6a).

Female. Concolorous with male; microtomentum of
parafrontal and parafacial slightly more brownish yellow than in
male; dorsum of thorax slightly more microtomentose, weakly
demarcating three shining stripes, but to the unaided eye scutum
only slightly less shining than in male.

Frons 1/4 width of head (0.25x; 0.23-0.27) and 3.275 times
ocellar span (3.00-3.54), widening slightly anteriorly; frontal vitta
with a few short and weak hairs, never with rows of bristlelike
hairs; triangular preocellar area often rugose, usually polished

P. shannoni—-_ 205

black but sometimes brownish microtomentose and subshining;
parafacial narrow as in male, its width equal to or barely wider
than 3rd antennal segment, about half width of frons (0.495x;
0.44-0.54), 1.62 times ocellar span (1.43-1.78), and 0.81 times
vibrissal interval (0.72-0.88); height of cheek 0.39 times
maximum length of eye (0.37-0.41).

Apparent normal length of adults, 8.5-9.5 mm.

First instar. Length up to 3 mm. Prothoracic fringe 14 wu
(6-23)/50. Cephalopharyngeal skeleton 240 wb (255-260)/7 in
length. Posterior region: stigmatal plates 26 w (21-29)/12 in
diameter; peritreme absent; mesostigmatal spines absent;
hyperstigmatal area with numerous spines, 6 mw (2-10)/50;
hypostigmatal spines scarce, as tubercles, 2 uw (1-4)/50; anal lobes
small. Dorsal cuticle with numerous spines, 8 ps (4-14)/50.
Ventral cuticle: spines of anterior patch typically three- to
four-pronged, 4 w (2-6)/50; medial band present, spines 6 wu
(2-12)/50; posterior band present.

Second instar. Prothoracic fringe 41 bm (24-62)/19.
Cephalopharyngeal skeleton 500 wu (405-530)/16 in length.
Posterior region: stigmatal plates 77 ms (69-92)/19 in diameter;
button indistinct; mesostigmatal spines scarce; hyperstigmatal
spines 10 w (6-16)/50; hypostigmatal spines numerous (Cf. 3rd
instar), as tubercles, 3 u (2-5)/50. Dorsal cuticle with numerous
spines, 12 ws (8-19)/50. Ventral cuticle: spines of anterior patch
7 pb (4-10)/50; medial band pronounced, spines 9 pw (6-14)/50;
posterior band pronounced.

Third instar. Peritreme usually closed; spines small and
scarce between stigmatal plates and on hypostigmatal area.
Prothoracic fringe 67 tt (41-82)/75. Cephalopharyngeal skeleton
780 w (580-940)/123 in length.

Puparium. Length 7.4 mm (5.5-9.4)/156; breadth 3.5 mm
(2.6-4.2)/171; thick-walled, shiny, black-brown. Posterior region
(Figs. 59a,b,; 60a,b): stigmatal plates 160 w (130-195)/364 in
diameter; distance between buttons 550 wu (375-730)/180, and
across stigmatal plates 1200 uw (850-1420)/180; stigmatal ratio
0.46 (0.43-0.52)/180; stigmatal area outlined in _ plaques,
frequently bisected; mesostigmatal spines small and _ scarce;
mesostigmatal plaques present, mesostigmatal folds infrequent;
hyperstigmatal spines usually as spines, 11 ws (4-24)/130,
sometimes as tubercles; hypostigmatal area variable, folds
normally present but sometimes absent; hypostigmatal spines as
tubercles, appearing as fine sandpaper, scarce, 6 w (2-10)/130;
circumstigmatal folds highly variable, normally rectangular with
pronounced folds (Figs. 59a, 60a), frequently with folds

206 Protocalliphora

surrounding stigmatal area (Figs. 59b, 60b), or, rarely, absent.
Dorsal cuticle (Figs. 59c, 60c) with numerous spines, 18
(6-41)/130; cuticular ridges pronounced (eastern forms, Fig. 59c)
or weak (western forms, Fig. 60c). Ventral cuticle (Figs. 59d,
60d); ventral band ratio 0.54 (0.51-0.67)/100; spines of anterior
patch 11 pw (6-27)/130; medial band pronounced, spines 14 pw
(6-27)/130; posterior band weak but present on all segments.
The ventral cuticle of the eastern forms (Fig. 59d) has
pronounced cuticular ridges while that of the western forms (Fig.
60d) has weak cuticular ridges.

Type Series. Holotype male, allotype, and 12 male, 12 female
paratypes, Algonquin Park, Ont., 1955 (G. F. Bennett), nest of
American robin. In the U.S. National Museum of Natural
History.

Additional paratypes: Canada [chiefly CNC, USNM]: 95
males, 96 females; Nova Scotia: Cariboo. Ontario: Algonquin
Park (nests of American robin, common grackle, gray catbird,
red-winged blackbird), London (nests of American robin,
red-winged blackbird) [U. Western Ontario]. Quebec: 4 mi N
Eardley, Laurentide Park, La Verendrye Provincial Park. Yukon
Territory: Dawson. Major series, 57 males, 53 females,
Algonquin Park, Ont. (Bennett), chiefly from nests of American
robin.

Contiguous United States: 150 males, 138 females:
Connecticut: Colebrook [Boston U.]. Delaware: Newark [U. Del.,
USNM]. Maine: Amherst and Aurora in Hancock Co., Dryden,
Enfield in Penobscot Co. [U. Maine, Field Mus. Nat. Hist.].
Massachusetts: Blue Hills Reservation and Groton [MCZ],
Concord and Martha’s Vineyard [Boston U.]. Montana: Hamilton
[Wash. State U., USNM]. New Hampshire: Ashland and
Peterboro [Boston U.], White Mts. [USNM]. New York: Babylon
[CAS], Granby Center in Oswego Co., Ithaca [Cornell U.]. Ohio:
Burton Township in Geauga Co. [Ohio State U.]. Pennsylvania:
Butler [Preston Colln.], Pittsburg [USNM, Mehner Colln.]. Rhode
Island: Buttonwoods in Kent Co. [Boston U.]. Utah: Franklin
Basin and Malibu Campground in Cache Co. [USNM, Whitworth
Colln.].

Non-Paratype Material: 140 males, 154 females. Many
specimens are in poor condition and have not been included in
the type series, although they can be recognized for additional
locality and host records. British Columbia: Aleza Lake and
Trinity Valley just E of Vernon, latter in nest of a junco.
Ontario: Algonquin Park in nests of white-throated sparrow and
hermit thrush, Dunrobin, Petawawa. Prince Edward Island:

P. shannoni = 207

Souris, in nest of common grackle. Maine: Levant, in American
tree swallow nest; Stetson in Penobscot Co. Maryland: Bethesda,
in nest of American robin. Michigan: Channing in Dickinson
Co. New York: Ballston Spa, Farmingdale, Yonkers.

Specimens Examined. 398 males, 401 females.

Distribution. Chiefly southeastern Canada (Ontario to Nova
Scotia) and northeastern United States (Michigan to Maine, south
to Ohio and Maryland), plus scattered records from British
Columbia and Yukon Territory, Montana, and Utah.

Basis of description of immature stages (all Algonquin Park,
Ont., some from type series): Larvae, Ist instar (7 specimens):
red-winged blackbird, July 1954 (3); common grackle, June,
1955 (4). Second instar (16 specimens): red-winged blackbird,
June, 1954 (3); American robin, August, 1954 (8); common
grackle, June, 1955 (5). Third instar (10 specimens): red-winged
blackbird, June, 1954 (4); American robin, June, 1955 (6).
Puparium (180 specimens): common grackle, June, 1952 (17) and
June, 1954 (27); red- winged blackbird, June 1954 (11); American
robin, August, 1954 (23), June, 1955 (67) and August, 1955 (35).

Hosts. Red-winged blackbird, eastern bluebird, gray catbird,
black-capped chickadee, common grackle, a junco, American
robin, white-throated sparrow, American tree swallow, brown
thrasher, hermit and wood thrushes, and house wren, probably
brown-headed cowbird (one of three fledglings from infested
wood thrush nest). Also, from puparia only, American
goldfinch, blue jay, gray [Canada] jay, eastern kingbird, song
sparrow, myrtle warbler, and cedar waxwing.

Ecology and Biology. Primarily in nests at intermediate or
canopy levels in forests. It has been found especially frequently
in robin nests, although this may in part reflect the facts that
robin nests are relatively common and easily observed and
sampled.

Variation. A long series from Hamilton, Mont., June 1951
(R. B. Spurrier) from robins’ nests showed variation in the
number of posterior bristles on the fore tibia. Of those which
could be counted (i.e., both fore legs present), 9 males and 10
females had one posterior bristle on both sides, one male and two
females had two on each side, and six males had two bristles on
one side only, left or right.

Remarks. P. shannoni is easily confused with sialia,
especially with males of the latter having relatively narrow frons.
Undoubtedly some past records of sialia are misidentifications,
especially of females. P. shannoni falls in a group of
intermediate species with few or no outstanding features;

208 = Protocalliphora

indeed its most distinctive character can be said to be the absence
of distinctive features in both sexes. However, the combination
of its narrow frons with the digitate surstylus will distinguish it
from others in the group. P. sialia has broader parafacials and
usually a broader frons. Males of aenea could easily be confused,
but the yellowish calypteres of male aenea will be a clue, and any
associated females of aenea will assure one beyond a doubt.
Brownish calypteres will separate some western species with
equally narrow frons and parafacials.

Females of shannoni are even less distinctive than males, but
the combination of polished or shining preocellar area, narrow
parafacials, and white calypteres will separate them from other
females of the sialia complex. If shannoni and sialia are mixed,
as they sometimes were in the study at Algonquin Park, the
obvious difference in parafacial width in both sexes will separate
the two reasonably well, aided by the more densely haired
parafacials of sialia. Associated puparia will also be definitive;
the densely long-spined and ridged puparium of sialia, with its
unusually long prothoracic fringe, is outstanding among Nearctic
Protocalliphora.

Puparia of shannoni are similar to those of deceptor and
metallica but can be separated by the presence of spines, rather
than tubercles, in the hyperstigmatal area. Protocalliphora
shannoni occurs across the continent and some differences
between the eastern (Fig. 59) and western (Fig. 60) forms can be
recognized. Eastern forms have rather pronounced cuticular
ridges while the western forms have, at best, weak cuticular
ridges.

Etymology. The specific name is in the genitive case,
honoring R. C. Shannon, calliphorid specialist whose seminal
work with Irene Dobroscky on the North American species of
Protocalliphora marked the first real attention to the rich North
American fauna in the genus.

Protocalliphora (P.) sialia Shannon and Dobroscky
(Figs. 15, 37, 61, Map 13)

Protocalliphora chrysorrhoea (Meigen).--Henshaw, 1908: 87-8
[Mass.; in type series of sialia], Miller, 1909: 3-7 [Penn.];
Banks, 1912: 21, pl. 3 [Mass., Penn.; description of larva].

Phormia chrysorrhea [sic].--Lloyd, 1922: 116 [Que.].

P. sialia 209

Protocalliphora splendida var. sialia Shannon and Dobroscky,
1924: 249, 251-2 [Widespread United States, especially
northeastern, and Ontario].

P. splendida form sialis [sic].--Johnson, 1925a: 216 [New
England; mixture of sialia, shannoni, metallica]; Johannsen,
1928: 827 [N.Y.; part metallica, part not verified].

P. splendida form sialia.--Johnson, 1927a: 1-3 [Mass.]; Johnson,
1927b: 33 [Mass.]; Johnson, 1927c: 77-9 [Mass.]; Johnson,
1927d: 204 [Maine, not verified].

P. splendida var. sialia.--Johnson, 1929a: 29-30 [Mass.].

P. splendida.--Lloyd, 1927: 111 [Correction of identification in
Lloyd, 1922]; McAtee, 1929: 105-111, in part [Md.].

P. sialia.--Johnson, 1920: 169-170 [Mass.]; Johnson, 1931:21-4
[Review of previous records]; Johnson, 1932: 26-8 [Mass.,
Vt., Ohio]; Wetherbee, 1932: 73-4 [Mass., Conn.].

P. splendida.--Mason, 1944: 232-247 [Mass.; mixture of sialia,
shannoni, metallica].

Apaulina sialia.--Hall, 1948: 201 [Northeastern U.S., s. Canada].

Protocalliphora spp.--Dodge and Seago, 1954: 53-4, in part [Ga.].

P. sialia.--Kenaga, 1961: 91-3 [Mich]; Boyes, 1961: 557
[Cytology]; Boyes and van Brink, 1965: 543 [Cytology]; Hall,
1965: 926 [Catalog, widespread]; Zeleny, 1970: 12 [Damage,
control]; Barbara Ann, 1970: 85-6 [Md.]; Boyes and Shewell,
1975: 442, 446 [Cytology]; Whitworth, 1976: 16ff. [Utah];
Zeleny, 1976: 123 [Biology, effect on birds]; Whitworth, 1977:
4933-B

Diagnosis. Sexes concolorous, monochromatic, shining dark
blue to bluish purple; male frons of medium width; female frons
typically with polished rugose preocellar area.

Male. Head black in ground color; parafrontal and parafacial
silvery microtomentose. Thorax shining metallic dark blue to
bluish purple, dorsum thinly microtomentose, viewed from
behind the microtomentum demarcating three broad shining
stripes, microtomentum especially conspicuous anterior to
transverse suture. Abdomen concolorous with thorax, but not as
dark, appearing more shining, tergites 2-4 with microtomentum
most evident when viewed at a low angle from behind.
Calypteres whitish.

Frons (Fig. 37) 0.09 times head width (0.08-0.11) and only
1.37 times ocellar span (1.13-1.67); frontal vitta narrow
posteriorly, widening anteriorly, at narrowest easily twice width
of an adjacent parafrontal; each parafrontal distinct throughout
but greatly narrowed above, the row of hairs on upper portion

210 =‘ Protocalliphora

almost merged with the row of frontal bristles, below becoming
two irregular rows and merging into the numerous hairs on the
parafacial; parafacial width approximately 1.5 times width of 3rd
antennal segment and 1.43 times width of frons (1.20-1.62),
nearly twice ocellar span (1.96x; 1.69-2.18), and about equal to
vibrissal interval (1.04x; 0.93-1.13); cheek height 0.42 times
maximum length of eye (0.38-0.45).

Thorax typically without accessory notopleural bristles and
without hairs on tympanic membrane and postalar wall. Fore
tibia with one posterior bristle.

Terminalia: cerci distally acuminate, parallel (as in Fig. 5b);
surstylus digitate, slightly wider basally, beyond the sides
subparallel, apex rounded (Fig. 15); aedeagus as usual in the
genus (cf. Fig. 6a).

Female. Concolorous with male, but thorax more heavily
microtomentose, viewed from behind the microtomentum and
three shining stripes evident to unaided eye; parafrontal and
parafacial grayish to light brown microtomentose, commonly with
a spot of changeable reflections on upper surface of parafacial,
opposite lunule, viewed from above the spot appearing dark
brown against the gray or light brown of parafacial and
parafrontal, from below bright yellowish gray against a black
background.

Frons at vertex 0.265 times width of head (0.24-0.29),
widening anteriorly and 3.31 times ocellar span (3.06-3.73);
frontal vitta parallel sided, usually with two rows of bristlelike
setae, each just mesad of a row of frontal bristles, rarely only a
few; preocellar area polished black, rugose, rarely dull and
microtomentose or the polished area greatly reduced in size;
parafacial moderately broad, width 0.555 times that of frons
(0.52-0.62), 1.84 times ocellar span (1.65-2.00), and 1.12 times
vibrissal interval (1.00-1.32); cheek height 0.44 times maximum
length of eye (0.42-0.46).

Apparent normal length of adults, 8.5-9.5 mm.

Egg. Length 1130 wu (1020-1205)/25; breadth 380 pw
(275-455)/25; cuticle white in color and ornamented with minute
spines.

First instar. Length up to 4 mm. Prothoracic fringe 13 wu
(8-21)/50. Cephalopharyngeal skeleton 220 w (205-240)/6 in
length. Posterior region: stigmatal plates 19 ww (16-25)/6 in
diameter; peritreme absent, mesostigmatal spines absent;
hyperstigmatal spines numerous, 6 bu (2-8)/50; hypostigmatal
spines scarce, 4 tt (2-6)/50; anal lobes large. Dorsal cuticle with
numerous spines, 8 mw (4-12)/50. Ventral cuticle: spines of

P. sialia 211

anterior patch typically three- to four-pronged, 3 (1-7)/50;
medial band present, spines6 (4-12)/50; posterior band present.

Second instar. Prothoracic fringe 52 (36-76)/50.
Cephalopharyngeal skeleton 525 (475-550)/13 in length.
Posterior region: button distinct; mesostigmatal spines scarce;
hyperstigmatal spines numerous, 14 (8-26)/50; hypostigmatal
spines numerous, 6 (2-12)/50. Dorsal cuticle with numerous
spines, 16 (8-31)/50. Ventral cuticle: spines of anterior patch
7 (4-10)/50; medial band pronounced, spines 11 (6-16)/50;
posterior band pronounced.

Third instar. Peritreme highly variable, open to closed;
meso- and hypostigmatal spines present. Prothoracic fringe long,
150 (80-200)/150. Cephalopharyngeal_ skeleton 975
(675-1100)/105 in length.

Puparium. Length 8.5 mm (5.8-10.8)/100; breadth 3.8 mm
(2.2-5.2)/100; thick-walled, dull, black brown. Posterior region
(Fig. 6la,b): stigmatal plates 190 (140-235)/250 in diameter;
distance between buttons 450 (235-690)/128, and across
stigmatal plate 1000 (655-1390)/128; stigmatal ratio 0.46
(0.43-0.56)128 (in parastigmate forms, i.e., poorly fed, the
stigmatal ratio ranges from 0.36 to 0.44); stigmatal area not
usually outlined in plaques; mesostigmatal spines scarce and
small; mesostigmatal folds and/or plaques present; hyperstigmatal
spines numerous, 35 (20-56)/150; hypostigmatal area with
numerous, irregular cuticular folds; hypostigmatal spines
numerous, 12 (4-22)/150; circumstigmatal folds present,
variable, folds normally pronounced and irregular (Fig. 61a,b)
but sometimes few and confined to hypostigmatal area and absent
in parastigmate forms. Dorsal cuticle (Fig. 61c) with numerous
spines, 34 (20-56)/150; cuticular ridges numerous and
prominent, frequently with two rows of spines per ridge, one
directed posteriorly and one anteriorly. Ventral cuticle (Fig. 61d):
ventral band ratio 0.89 (0.80-1.0)/100; spines of anterior patch 18

(6-33)/150; medial band entire and pronounced, spines 27
(16-45)/150; posterior band pronounced; ventral cuticle with
pronounced cuticular ridges.

Puparia formed by larvae which have fed insufficiently are
frequently found in infestations of sialia. These puparia are
shorter, light brown in color and thin-walled; the stigmatal plates
are usually close together, giving a stigmatal ratio of 0.37
(0.36-0.44) - the parastigmate forms. The spines on the dorsal
and ventral cuticle are crowded, giving the impression of more
than the usual number of spines; the three ventral spine bands are
frequently elided.

212 ~~ Protocalliphora

Type Series. For some reason, no holotype was designated by
the authors, although they did so for all other new taxa
(including new varieties) described in their 1924 revision. Hall
(1948) recorded the failure to establish a holotype, but did not
designate a lectotype. Shannon and Dobroscky specified that
sialia was based only on females, and detailed records were given
for a number of specimens. These females, being the material on
which the variety was based, constitute the syntype series of
sialia, in the absence of any contrary or positive statement by the
authors. Many of these have been located in collections, a
lectotype has been selected, and lectoparatypes labeled. A few
of these original females must be referred to other species.

Lectotype and two paralectotypes, female, Shawnee, on
Delaware River, Penn., July 31, 1908, "on young bluebird," and
labeled "Protocalliphora splendida > sialia S. & D." by Shannon
himself. Published by Miller (1909) as P. chrysorrhoea.

Paralectotypes, all females [USNM except as noted]: Two,
Wellesley, Mass., one labeled Aug. 10, 1907 (Mrs. E. F. Everett);
one, Blue Hill, Mass. (N. Banks) [MCZ]; one, "Ottawa, Ont.", July
23, 1922 (H. Lloyd), "parasitic on nestling bluebird" (apparently
one of the specimens published by Lloyd, 1922, as Phormia
chrysorrhoea from 5 mi W of Hull, Quebec).

Syntypes of sialia referred elsewhere [all USNM]: two,
Alaska: Savonoski, Naknek Lake (J. S. Hine), paratypes of
Apaulina sapphira Hall (1948); one, New Hampshire: White Mts.
[H. K. Morrison] to shannoni n. sp.; one, New Mexico: top of Las
Vegas Range, 11000 ft, end of June (T. D. A. Cockerell) to
spatulata n. sp. (probable); many, Virginia: East Falls Church (E.
A. Chapin), and one, Rosslyn, May 29, 1913 (R. C. Shannon),
brown thrasher, to metallica; two, Washington: Almota (A. L.
Melander) to hirundo.

Specimens Examined. 798 males, 746 females, many
unassociated puparia.

Distribution. Widespread, Alaska and Northwest Territories
to New Brunswick, south to California, New Mexico, Iowa, and
Virginia, also northern Georgia, Mexico (Durango, Puebla).

Alaska: Eagle, Kasilov and Sterling on Kenai Peninsula,
Palmer.

Canada: British Columbia: Haney, Kamloops, Penticton,
Quesnel, Rike Creek in Becher’s Prairie. New Brunswick: Acadia
Forestry Station, Chamcook, Komhibouguac National Park.
Northwest Territories: Norman Wells in District of Mackenzie.
Ontario: Algonquin Park (143 males, 178 females, many puparia),
Elgin, Guelph, Mallorytown Landing, Marmora, Ogoki, Osgoode,

P.sialia 213

Ottawa, Toronto. Quebec: Barriére St. Anne in Laurentides Park,
La Trappe, La Verendrye Provincial Park, Masham Township in
Gatineau Co., Montreal, Parc du Mont Tremblant, St. Pierre de
Wakefield. Yukon Territory: Kluane Lake.

Contiguous United States, East: Connecticut: Pamfret,
Stamford, Waterbury. Delaware: Yorklyn. Georgia: Mt. Enotah
in Towns and Union Counties, Rabun Bald in Rabun Co. Iowa:
Cedar Falls. Maine: Aurora, Enfield, Levant, Mariaville.
Maryland: Bell, Beltsville, Bowie, Catonsville, Laurel.
Massachusetts: Babson Park, Belmont, Blue Hill, Dover, Groton,
Needham, Rock, Waban, Wellesley, West Bridgewater, Worcester.
Michigan: Ann Arbor, Cheboygan Co., East Lansing, Midland,
Vicksburg, "Upper Peninsula" (no locality). New York: Ballston
Spa, Mt. Hurricane in Essex Co., Voorheesville in Albany Co.
Ohio: Columbus, Leetonia in Columbiana Co., Oak Openings
Park in W Lucas Co., Toledo. Pennsylvania: Butler, Clarksville,
Kennett Square near Philadelphia, Shawnee-on-Delaware. South
Dakota: No locality. Virginia: mouth of Aquia Creek in Stafford
Co., East Falls Church, Falls Church, Front Royal, Hopewell.
West Virginia: Ona.

Contiguous United States, West: Arizona: Chiricahua Mts.,
Portal, Ramsey Canyon in Huachuca Mts. California: Fredonyer
Peak in Lassen Co. (25 mi N Susanville), Carmel Valley in
Monterey Co., French Camp in San Joaquin Co., O’Neals in
Madera Co., Roney Flat in Modoc Co., Sagehen near Hobart
Mills in Lassen Co. Colorado: Colorado Springs, Ft. Collins, Weld
County. Idaho: Moscow Mt. Montana: Hamilton, top of Mt.
Sentinel in Missoula Co., Ravalli Co. New Mexico: Tomay in
Valencia Co. Oregon: Klamath Forest National Wildlife Refuge
in Klamath Co. Utah: Benson, Mendon, Newton, Smithfield and
Wellsville in Cache Co., Moroni, near Roosevelt in Duchesne Co.,
Snowville in Box Elder Co. Washington: Dartford, Kamiak Butte
near Pullman, Pullman, Seattle. Wyoming: La Barge in Lincoln
Co.

Mexico: Male, 11 m W Durango, 7000 ft, June 20, 1964 (J. F.
McAlpine) [CNC]. Puebla: 3 males, San Martin, Texmelucan,
May 26, 1922 (E. G. Smyth) [USNM].

Basis of description of immature stages (all material from
Algonquin Park, Ont.): Egg: from caged female, July 1955.
Larvae, Ist instar (6 specimens): common grackle, June, 1953.
Second instar (13 specimens): common grackle, June, 1955.
Third instar (10 specimens): common grackle, June, 1955 (6);
European starling, June, 1952 (2) and June, 1955 (2). Puparium
(150 specimens): common grackle, June, 1955 (59); European

214 ~~ Protocalliphora

starling, June, 1955 (10); rough-winged swallow, August, 1950
(6); downy woodpecker, July, 1955 (10); barn swallow, July, 1955
(65).

Hosts (Eastern): Eastern bluebird, yellow-shafted flicker
(puparia only), great crested flycatcher, common grackle, eastern
kingbird, purple martin, white-breasted nuthatch, eastern
screech-owl (puparia only), eastern phoebe (puparia only),
common raven, American robin, house and song sparrows,
European starling, swallows (bank, barn, cliff, rough-winged,
American tree), downy woodpecker, and house wren. Shannon
and Dobroscky (1924) recorded sialia from the common crow at
Ithaca, N.Y., but we have not seen such material; there might
have been confusion with females of avium.

Hosts (Western). Brewer’s blackbird, mountain and western
bluebirds, mountain chickadee, American crow, ash-throated
flycatcher, American kestrel [sparrow hawk], western kingbird,
white-breasted nuthatch, flammulated and great horned owls,
western screech-owl, Say’s phoebe, common raven, American
robin, house sparrow, European starling, various swallows (bank,
barn, cliff, American tree, and violet-green), and house wren.
P. sialia was also reared from a ringed turtle-dove nest in
Colorado by R. M. Stabler, but the hosts were caged birds in
experimental studies of Trichomonas and not a feral established
host population.

Part of the type series (syntypes) of sialia, from Ithaca, N.Y.,
was recorded as reared from "crows and robins." Many of the
specimens may actually have been part of the series from crows
herein recorded as avium. P. sialia occurs typically in cavity
nests, or in low-nesting birds, and the common crow would be a
most unusual host.

Ecology and Biology. This species occurs frequently in both
cavity and non-cavity nests in man-made structures, and in
cavity nests anywhere. The biology has been studied extensively
by Bennett in Algonquin Park, Ont., where it occurred commonly
in nests of common grackle and barn swallow.

Variation. The ratios given in the description are those of
sialia from the East, the typical range of the species. Specimens
from the western United States and southwestern Canada varied
slightly from the typical form and were observed closely to see
whether they might represent a western subspecies, or even a
closely related species, similar to the asiovora-avium pairing. No
consistent or significant differences could be found and all are
here referred to sialia. Ratios of eastern and western forms are

P.sialia 215

given here for comparison, based on 25 of each sex from each
region (Table IX).

Occasional specimens show some slight aberration, which we
have come to expect in these flies. One male from Mt. Enotah,
Ga., has accessory notopleural bristles like those of braueri, and
females from Kamloops, B.C., and Hamilton, Mont., have two
posterior bristles on one fore tibia but only one on the other, the
former having the two on the right tibia, the latter two on the
left.

Remarks. P. sialia is one of the commonest species of the
genus. It is frequently found in nests of swallows and the
common grackle, but it has also been recorded from a number of
other birds. The broad silvery parafacial of the male is
distinctive, as is the polished black preocellar area of the female,
at least those of the East. A polished black preocellar area is
found only in hesperioides and some shannoni, in addition to
sialia, so this is an especially useful characteristic:for females.
In the West females often have the _ preocellar area
microtomentose, but no other differences could be found. In
view of the distinctive puparium like that of eastern specimens,
with strong spines and especially long prothoracic fringe, the
western specimens are here recorded as sialia but with the
reservation that further attention should be given to the western
population.

Shannon and Dobroscky’s original series of sialia was greatly
mixed, since the name was applied to blue females, which
brought together under the one name several distinct species.

P. sialia and P. hirundo are similar, but the latter has
distinctly gray microtomentose preocellar area in both sexes,
whereas that area is shining black in both sexes in sialia.
Separation of females of western sialia and hirundo will be
difficult or impossible, however, unless there are associated males
or puparia. Likewise, unassociated females of avium and sialia
will be difficult to distinguish, although avium has slightly
broader parafacials that are usually slightly convex and appearing
bulging. P. avium is also usually larger and more robust.

P. sialia can also be confused with the new species shannoni,
especially since the two occupy similar ranges in the East. The
wider frons of the male and wider parafacial in both sexes are
characteristic of sialia, but teneral males of sialia may have a
narrow frons that is suggestive of shannoni.

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P. spatulata 217

Puparia of sialia are similar to those of avium, chrysorrhoea,
and hirundo. They can be separated from these and all other
Protocalliphora by the unusually long prothoracic fringe.

One doubtful record should be mentioned because of the
myiasis involved. Lincoln (1931) recorded "spendida" larvae
from a barn swallow nestling at Fairdale, N. Dak. The maggots
were clinging to its eyelids and throat, and some in the nostrils.
Unfortunately these maggots are very small 3rd-stage larvae and
one can only say that they are probably sialia, but possibly
hirundo.

Protocalliphora (P.) spatulata, new species
(Fig. 27, 62, Map 12)

Protocalliphora "metallica."--James, 1955: 25 [Calif].

"Blowfly larvae," nest of horned lark.--Verbeek, 1967: 216
[Wyo.].

"Blowfly larvae," nest of water pipit.-- Verbeek, 1970: 440 [Wyo.;
larvae only, apparently P. spatulata].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic dark blue to bluish purple; calypteres opaque white;
frons of male relatively broad; parafacials broad in both sexes;
male with surstyli in‘ profile narrow, parallel sided, curved,
becoming spatulate distally (Fig. 27).

Male. Head dark, entirely black or nearly so, parafrontal and
parafacial obscurely brownish microtomentose, microtomentum
on parafrontals thinning posteriorly to the chiefly polished black
prevertical area. Thorax shining, dark metallic blue to bluish
purple, scutum thinly dark gray microtomentose, especially
anteriorly, but shining stripes not clearly delineated and scutum
appears entirely dark and shining, especially to unaided eye, and
weakly microtomentose only when viewed from_ behind.
Abdomen approximately concolorous with thorax, but usually a
brighter dark metallic blue to bluish purple, appearing shining in
direct view but thinly dark gray microtomentose when viewed
from behind at a low angle, Sth tergite shining. Calypteres
Opaque white, outer rims tinged with yellow.

Frons relatively broad, 0.12 times head width (0.10-0.13) and
obviously wider than ocellar span (1.60x; 1.33-1.79); parafrontal
with hairs outside the frontal bristles on its full length at least to
level of ocellar bristles, parafrontal not narrowed above but
continued of same width to vertex; prevertical area smooth and

218 Protocalliphora

polished, bare of microtomentum, rarely thinly microtomentose
about bases of bristles and hairs or across the middle; preocellar
area narrow, elongate and acuminate, rugose and subshining;
parafacial broad, obviously much wider than 3rd antennal
segment, its width slightly greater than the broad frons (1.19x;
1.08-1.38), nearly twice ocellar span (1.90x; 1.71-2.29), and 0.90
times vibrissal interval (0.79-1.09), with numerous coarse black
hairs in four irregular rows; cheek broad, its height nearly half
(0.47) maximum length of eye (0.42-0.49).

Thorax without accessory notopleural bristles and without
hairs on tympanic membrane, a few inconspicuous hairs
sometimes present on postalar wall. Fore tibia with one posterior
bristle.

Terminalia: cerci distally acuminate, parallel (as in Fig. 5b);
surstylus (Fig. 27) moderately narrow, parallel-sided for most of
length, curved, usually widening rather abruptly at distal end to
appear spatulate; aedeagus as usual in the genus (cf. Fig. 6a).

Female. Concolorous with male; parafrontal and parafacial
decidedly brownish microtomentose; dorsum of thorax slightly
more microtomentose than in male, but microtomentum still not
delineating the three shining stripes usually evident in females of
the genus, the thorax appearing to unaided eye as entirely shining
and unstriped.

Frons broad, nearly twice width of eye and nearly 1/3 width
of head (0.31x; 0.29-0.33) and 3.56 times ocellar span (3.10-4.00),
parallel sided but gradually widening anteriorly; frontal vitta
with a number of coarse black bristlelike and mesoclinate setae;
preocellar area sometimes subshining but always evenly brownish
microtomentose; parafacial broad, as in male, its width nearly
half as broad as head (0.485x; 0.42-0.54), 1.73 times ocellar span
(1.40-2.00), and 0.98 times vibrissal interval (0.85-1.10); height
of cheek about half maximum length of eye (0.50x; 0.45-0.54).

Apparent normal length of adults, 9-10 mm.

Third instar. Peritreme closed; spines sparse between
stigmatal plates and in hypostigmatal area. Prothoracic fringe 58
lL (34-90)/100 (measurements from both larvae and puparia).
Cephalopharyngeal skeleton 715 wu (670-840)/16 in length.

Puparium. Length 8.1 mm (7.2-8.8)/16; breadth 3.6 mm
(3.1-3.9)/16; medium thick-walled, dull, dark brown. Posterior
region (Fig. 62a): stigmatal plates 178 wu (150-200)/32 in
diameter; distance between buttons 434 mw (320-500)/16, and
across stigmatal plates 1010 wu (910-1230)/16; stigmatal ratio 0.43
(0.34-0.51)/16; stigmatal area outlined in plaques and weak folds;
mesostigmatal spines virtually absent; mesostigmatal folds and

P. spatulata 219

plaques pronounced, bisecting the stigmatal region;
hyperstigmatal spines rather slender, 30 u (17-47)/160 in length;
hypostigmatal area with numerous, irregular, pronounced folds;
hypostigmatal spines rather sparse, 20 u (8-43)/160 in length,
most spines short but a few long ones on each puparium;
circumstigmatal folds concentric, rather weak. Dorsal cuticle
(Fig. 62b) with numerous spines, shortest towards the center line,
28 uw (14-46)/160; cuticular ridges usually weak. Ventral cuticle
(Fig. 62c): ventral band ratio 0.83 (0.72-0.92)/16; spines of
anterior patch 17 w (8-29)/160, rather slender; medial band
pronounced, frequently almost contiguous with anterior band,
spines slender, 24 pu (11-38)/160; posterior band pronounced.
Type series. Holotype male, allotype, and 26 male, 25 female
paratypes, Two Ocean Mt., near Togwotee Pass, Teton Co., Wyo.
(N. R. French), parasite on rosy finch, larvae collected July 29,
1951, pupated about middle of August, and adults emerged Sept.
9, 1951. In the U. S. National Museum of Natural History
paratypes in French Collection, U. Colo., CNC, CAS, and Amer.
Mus. Nat. Hist., by courtesy of the collector. Additional
paratypes: Alaska: male, female, Sheenjek River, NE Alaska, 68°
26’-32’ N, 143° 45°-55’ W, June 2, 1956 (G. B. Schaller)
[USNM]; male, Crowbill Mountain, Cape Thompson [N of
Kotzebue], June 7, 1960 (W. C. Hanson) [Wash. State U.].
Canada: Northwest Territories: male, Tununuk [Mackenzie
District], Aug. 15, 1930 (O. Bryant) [USNM]; male, Wrigley [On
Mackenzie River north of Ft. Simpson], June 7, 1969 (G. E.
Shewell) [CNC]. Ontario: male, female, Ogoki [At juncture of
Ogoki and Albany Rivers, NE of Lake Nipigmy, N of Lake
Superior] [CNC]. Yukon Territory: male, Mile 87, Dempster
Highway, Aug. 4-8, 1973 (G. and D. M. Wood) [CNC].
Contiguous United States: California [chiefly U. Calif,
Berkeley]: male, Lassen Peak, 7500 ft, Shasta Co., July 18, 1949
(E. L. Atkinson); male, Sentinel Dome, Yosemite, 8117 ft, June
22, 1931 ["metallica" in James 1955]; male, Yosemite, 3880-4000
ft, June 22, 1931; male, Sonora Pass, 9-10000 ft, Mono Co., July
11, 1957 (J. W. MacSwain): male, Sonora Pass, Tuolumne Co.,
June 26, 1960 (J. F. Lawrence) [CAS]. Colorado: female,
Doolittle Ranch, 9800 ft, Mt. Evans, Aug. 3, 1961 (J. G.
Chillcott) [CNC]. Montana: 2 males, top of Squaw Peak, 7996 ft,
near Missoula, July 14, 1951 and July 28, 1952 (J. A. Chapman)
[USNM]; male, West Fork, Ravalli Co., June 10, 1934 (W. L.
Jellison) [USNM]. New Mexico: female, top of Las Vegas Range
[Lower end of Sangre de Cristo Mts., NW of Las Vegas], 11000
ft, end of June (T. D. A. Cockerell) [USNM] [paratype of sialia].

220 ~~ Protocalliphora

Wyoming: male, Riverside, Yellowstone Park, Aug. 4, 1918 (A.
L. Melander) [USNM]; 5 males, Park Co., 10500 ft, above
timberline on Beartooth Plateau, about 1 km N of Beartooth Pass,
larvae collected Aug. 20, 1964, nest of horned lark (N. A. M.
Verbeek), and 4 males, 5 females, same data except larvae
collected July 20, 1964 [USNM].

Non-paratype Material: male, Summit Mt. Moscow, Idaho,
July 1, 1932 (J. M. Aldrich) [USNM]; 2 females, Park Co., Wyo..,
larvae collected Aug. 20, 1964 (Verbeek) [USNM]; male, Yukon
Territory, Mile 87, Dempster Highway, Aug. 4-8, 1973 (G. and
D. M. Wood) [CNC]. Also, 12 larvae from the same Park Co.,
Wyo., locality, collected July 29, 1963 (Verbeek) from nest of a
water pipit [USNM], appear to be spatulata.

Specimens Examined: 53 males, 37 females.

Distribution: Western, apparently chiefly at high elevations,
typically in rocky alpine tundra, Alaska to western Ontario, south
to California and New Mexico,

Basis of description of immature stages: Larva, 3rd instar (4
specimens): Park Co., Wyo., horned lark (Verbeek, No. XV).
Puparium (16 specimens): Park Co., Wyo., horned lark (Verbeek
No. VII) (9) and (Verbeek No. XII) (7).

Hosts: Rosy finch, horned lark, water pipit (probable, larvae
only).

Ecology and Biology: The known hosts, two and probably
three, are ground-nesting birds, characteristically in rocky alpine
tundra. Most of the known specimens have been taken at high
elevations in the West.

The type series from Two Ocean Mt. was reared from a rosy
finch nest. Dr. French (personal communication) has supplied
the following information: "The parasites were first noticed on
July 29, 1951, when four young birds were recovered from the
nest hole for banding. At this time six larvae were seen clinging
to the bare areas of the lower breast and abdomen of two of the
young birds. On August 2, 1951, when the empty nest was
collected it was found to contain a large number of larvae." They
pupated about the middle of August and the first adult flies
appeared about September 9th.

Variation. Normally there is only one posterior bristle on the
fore tibia, but specimens have been seen in which either a right
or a left tibia had none or two, the other tibia one. Such
examples would confuse the user of a key, if he had only one
specimen, and there would be even more confusion if two or
none occurred on both fore tibiae. Luckily in males the

P. spenceri 221

relatively wide frons and parafacial and the characteristic surstyli
will overcome any difficulty with the variation.

The non-paratype male from Yukon Territory is a "runt",
obviously an underfed individual, only 6.5 mm in length
compared with the normal of 9-10 mm.

Remarks. P. spatulata is one of the group of species with
narrow curved surstylus. Its relatively broad frons, broad
parafacial, and broad and very setose parafrontal make it one of
the most distinctive species in that group. It is closest to
parorum, and the differences are not great.

The puparia of this species can be confused with chrysorrhoea
and to a lesser extent with avium and sialia. P. spatulata can be
best separated from the latter two by the lack of irregular
circumstigmatal folds and from the first by the longer
prothoracic fringe; it differs from all three in lacking cuticular
ridges.

Etymology. The specific name, an adjective referring to the
shape of the surstylus, is derived from the Latin spatula,
diminutive of spatha, a flat instrument for stirring.

Protocalliphora (P.) spenceri, new species
(Figs. 32, 63, Map 14)

Protocalliphora sp. XV.--Whitworth, 1976: 20, 33, 38 [Utah].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic blue to bluish purple, female somewhat more
microtomentose and slightly duller; calypteres white; male with
surstylus curved and strongly narrowed.

Male. Parafrontal and parafacial with fine shining
microtomentum, yellowish to light brownish tinted, not forming
a distinct changeable spot on upper part of parafacial. Body
shining, metallic blue to dark bluish purple or bluish black,
scutum very thinly microtomentose, delineating three shining
stripes anterior to transverse suture, the abdomen highly shining
but thinly microtomentose when viewed at a low angle from
behind, 5th tergite more shining. Calypteres opaque whitish,
outer rims faintly yellowish.

Frons narrow, 0.07 times head width (0.06-0.8/17) and
usually slightly wider than ocellar span (1.14x; 1.00-1.30/71);
preocellar area a polished line or narrow ridge, sometimes rugose;
lower parafrontal and parafacial sparsely to considerably haired,
in two or three irregular rows; width of parafacial about equal to

222 _Protocalliphora

breadth of 3rd antennal segment, slightly greater than width of
frons (1.48x; 1.29-1.67/17), 1.68 times ocellar span
(1.50-1.90/17), and 0.66 times vibrissal interval (0.57-0.79/17);
height of cheek 0.38 times maximum length of eye
(0.36-0.41/17); occiput with two irregular rows of black hairs
behind row of postocular setae.

Thorax without accessory notopleural bristles, tympanic hairs,
and hairs on postalar wall. Fore tibia with a single posterior
bristle.

Terminalia: cerci distally acuminate, parallel (as in Fig. 5b);
surstylus curved, strongly narrowed on distal portion (Fig. 32);
aedeagus as usual in the genus (cf. Fig. 6a).

Female. Concolorous with male, but gray microtomentum
denser on scutum and abdomen, the shining scutal stripes more
distinct.

Frons not as wide as in many females of Protocalliphora, 1/4
width of head (0.25x; 0.24-0.275/24) and 3.22 times ocellar span
(2.86-3.62/24); frontal vitta with small irregular polished spot
adjacent to median ocellus, sometimes microtomentose but
subshining; prevertical area dull, microtomentose like rest of
parafrontal; parafacial about equal to breadth of 3rd antennal
segment, 0.485 times width of frons (0.45-0.52/24), 1.56 times
ocellar span (1.31-1.69/24), and 0.75 times vibrissal interval
(0.68-0.85/24); height of cheek 0.40 times maximum length of
eye (0.37-0.435/24).

Apparent normal length of adults, 7.5-8 mm.

Third instar. No specimens examined, characters derived
from puparium. Prothoracic fringe 47 | (23-80)/80.
Cephalopharyngeal skeleton 860 w (770-935)/11 in length.

Puparium. Length 7.5 mm (6.7-8.3)/6; breadth 3.4 mm
(3.0-4.0)/15; medium thick-walled, dull, brown. Posterior region
(Fig. 63a): stigmatal plates 160 wu (135-190)/30 in diameter;
distance between buttons 475 w (380-540)/15, and across
stigmatal plates 1030 mw (870-1300)/15; stigmatal ratio 0.49
(0.46-0.53)/15; stigmatal area outlined in large cuticular plaques;
mesostigmatal area with pronounced plaques; mesostigmatal
spines present, but scarce and small; hyperstigmatal spines
numerous, stout, 21 pw (10-50)/150; hypostigmatal area with
reduced folds, sometimes pronounced; hypostigmatal spines
generally scarce, 9 &¢ (4-25)/150; circumstigmatal folds generally
absent or at best weak, concentric. Dorsal cuticle (Fig. 63b) with
numerous spines, 20 uw (10-50)/150; cuticular ridges weak.
Ventral cuticle (Fig. 63c): ventral band ratio 0.70 (0.40-0.80)/15;
anterior band pronounced with anterior patch distinct, spines 9

P. spenceri 223

wt (4-17)/150; medial band pronounced, spines 14 pt (6-30)/150;
posterior band pronounced, present on all segments, with
numerous spines.

Type Series. Holotype male, allotype, and 11 male, 8 female
paratypes, Okanagan Landing, B.C., July 19-28, 1941 (G. J.
Spencer), nest of yellow warbler collected July 12. Holotype and
allotype in the U.S. National Museum of Natural History, by
courtesy of the late Professor Spencer; paratypes in the Canadian
National Collection and the University of British Columbia.
Additional paratypes: 5 males, 17 females, Ft. St. James, B.C.,
American redstart nest collected July 16, 1947 (J. Grant), flies
reared in laboratory Aug. 10, 1947 (G. J. Spencer) [UBC and
USNM]: male, Waha, Idaho, Aug. 12, 1923 (A. L. Melander)
[USNM, labeled, but not published, as a paratype of cuprina}; 3
males, 3 females, Franklin Basin, Cache Co., Utah, July 27, 1971
(T. L. Whitworth), nest of warbling vireo [USNM and USU];
male, female, Ninette, Manitoba, May 5, 1958 (J. F. McAlpine),
"at bleeding stump Acer negundo" [CNC].

Non-paratype Material: 3 males, 1 female, Okanagan
Landing, B.C., from series from nest of yellow warbler; male,
female, Ft. St. James, B.C., from series from nest of American
redstart; Trinity Valley, B.C., puparia only, from nest of
slate-colored junco [UBC].

Specimens Examined. 26 males, 32 females.

Distribution. British Columbia, Manitoba, Idaho, Utah.

Basis of description of immature stages: Puparium (15
specimens): Trinity Valley, B.C., slate-colored junco, August,
1946 (3); Fort St. James, B.C., American redstart, July, 1947 (5);
Okanagan Landing, B.C., yellow warbler, July, 1941 (2); Franklin
Basin, Cache Co., Utah, July 1971, warbling vireo (5).

Hosts. American redstart, warbling vireo, yellow warbler;
also, from puparia only, slate-colored junco.

Ecology and Biology. Unknown.

Variation. An extremely aberrant variation was found among
the paratypes from Okanagan Landing, B.C. Two females lacked
reclinate fronto-orbital bristles, a characteristic considered to be
typical of and confined to the subgenus Trypocalliphora and its
single included species braueri. In fact, these two specimens
would cause difficulty in the very first couplet of the key to
females. One male and one female of the Cache Co., Utah series
have two posterior bristles on the fore tibia.

Remarks. The curved surstylus associates spenceri with a
certain group of species, at least a grouping of convenience.
Males of these species, except for one or two, are difficult, and

224 ~~ Protocalliphora

unassociated females are virtually impossible. For best
identification, the sexes must be associated and preserved with
puparia. The extremely narrow frons and parafacial will separate
spenceri males from the other species. P. bicolor and P. metallica
also have such a narrow frons but wider parafacials, and the
former is further distinguished by having the 5th tergite cupreous
in the male.

The main series from Okanagan Landing, B.C., was a mixed
infestation of spenceri and cuprina, but luckily both sexes of
these two species are easily distinguished. Males of the two have
distinct surstyli, and in females of cuprina the 5th tergite is
cupreous.

Puparia of spenceri are separated with difficulty from those
of halli and seminuda. Separation is based on a consideration of
all the characters.

Etymology. We take pleasure in dedicating this species to the
late Professor G. J. Spencer of the University of British
Columbia, whose long interest in Protocalliphora led to the
accumulation of fine series of adults and immature stages, which
he generously loaned to us for study.

Protocalliphora (P.) tundrae, new species
(Figs. 23, 36, 64, Map 12)

Calliphora azurea (Fallén).--Lundbeck, 1895: 114 [Hekla-Havn,
East Greenland; same record repeated in several subsequent
compilations on the insects of Greenland].

C. azurea.--Nielsen, 1907: 395 [Same record, with addition of

date of collection, June 11, 1892].

Protocalliphora sp.--Oliver, 1963: 178 [Ellesmere I., N.W.T.].

Diagnosis. Sexes concolorous, monochromatic, shining
metallic dark bluish; calypteres white; frons of male of medium
width, preocellar area broadly triangular, bright gray
microtomentose; surstylus narrow, curved.

Male. Head chiefly black in ground color, parafrontal and
parafacial silvery gray microtomentose. Body shining, metallic
dark bluish to bluish purple, thorax darker than abdomen,
scutum viewed from behind with two stripes of gray
microtomentum that demarcate a broad median shining stripe and
the broad lateral shining areas; abdomen brighter than scutum,
shining in direct view but gray microtomentose on tergites 2-4
when viewed from behind at a low angle. Calypteres white.

P.tundrae 225

Frons of moderate width (Fig. 36), at narrowest 0.135 times
head width (0.12-0.16) and 1.43 times ocellar span (1.22-1.73);
ocellar tubercle typically with 6-8 small bristles behind the strong
ocellar bristles; preocellar area broadly triangular, entirely and
evenly bright gray microtomentose; upper end of row of frontal
bristles usually opposite anterior end of preocellar triangle;
parafrontal narrowed just before vertex, silvery gray
microtomentose, parafrontal hairs in two irregular rows below,
narrowing to one row above and ending well before end of row
of frontal bristles; parafacial silvery gray microtomentose, with
numerous fine black hairs in three irregular rows, its width
obviously greater than breadth of 3rd antennal segment, equal to
width of frons (1.00x; 0.91-1.13), 1.43 times ocellar span
(1.22-1.62), and 0.86 times vibrissal interval (0.73-1.00); height
of cheek 0.46 times maximum length of eye (0.43-0.48); occiput
with 3-4 irregular rows of black hairs behind row of postocular
cilia.

Thorax lacking accessory notopleural bristles; tympanic
membrane and postalar wall with few short weak and
inconspicuous hairs. Fore tibia typically with two posterior
bristles.

Terminalia: cerci long and narrow to acute apices (cf. Fig.
5b); surstylus long and narrow, strongly curved (Fig. 23);
aedeagus as usual in the genus (cf. Fig. 6a).

Female. Concolorous with male, but slightly more gray
microtomentose. Frons nearly 1/3 width of head (0.31x;
0.30-0.32) and over three times ocellar span (3.16x; 2.89-3.62);
preocellar area entirely and evenly dull, heavily yellowish gray
to brownish gray microtomentose, as is microtomentum of
parafrontal and parafacial; parafacial with numerous fine black
hairs in 3-4 irregular rows, definite changeable spot along upper
margin, width nearly twice breadth of 3rd antennal segment, 1/2
breadth of frons (0.50x; 0.44-0.54), 1.59 times ocellar span
(1.29-1.75), and nearly equal to vibrissal interval (0.99x;
0.76-1.04); height of cheek nearly half maximum length of eye
(0.49x; 0.47-0.51).

Apparent normal length of adults, 7.5-8.5 mm. (The two
smallest specimens are only 5.5 mm).

Third instar. No specimens examined, characters derived
from puparium. Prothoracic fringe 54 pm (26-84)/75.
Cephalopharyngeal skeleton 860 wu (770-1000)/10 in length.

Puparium. Length 7.5 mm (6.8-7.8)/12; breadth 3.1 mm
(2.6-3.4)/12; thin-walled, dull, light brown. Posterior region
(Figs. 64a,b): stigmatal plates 180 wu (160-195)/23 in diameter;

226 ~—_- Protocalliphora

distance between buttons 565 pw (470-630)/12; and across
stigmatal plates 1080 wu (760-1190)/12; stigmatal ratio 0.51
(0.48-0.52)/12; stigmatal area somewhat depressed, faintly
outlined in small plaques; numerous mesostigmatal spines present;
mesostigmatal folds and plaques weak; hyperstigmatal spines
numerous, 31 wu (16-46)/160; hypostigmatal area with few
cuticular ridges or folds; hypostigmatal spines numerous, 23 pu
(8-42)/160; circumstigmatal folds weak and concentric. Dorsal
cuticle (Fig. 64c) with numerous, weak, cuticular ridges; spines
numerous, 29 uw (10-46)/160. Ventral cuticle (Fig. 64d): ventral
band ratio 0.57 (0.51-0.62)/10; spines of anterior patch slender,
17 pw (8-28)/150; medial band pronounced, spines 20 wu
(8-38)/160; posterior band reduced on anterior 5-6 segments and
vestigial or absent on segments six to twelve.

Type Series (all but a few reared and accompanied by
puparia). Holotype male, allotype, and 20 paratypes (8 males, 12
females), Northwest Territories: Ellesmere Island, Hazen Camp,
81° 49’ N, 71° 18° W, various dates of emergence July 20-Aug.
3, 1963 (D. R. Oliver), ex snow bunting nest. In the Canadian
National Collection, Ottawa; some paratypes in USNM.
Additional paratypes [CNC except for representatives in USNM
and one in Copenhagen Museum]: Canada, Northwest Territories:
Ellesmere I., same locality as holotype, 12 males, 20 females,
various dates, Aug. 2-19, 1963 (R. E. Leech), ex snow bunting
nest; 5 males, 5 females, same locality, various dates July
27-Aug. 1, 1962 (R. B. Madge), reared but host not given; one
female, same locality, June 19, 1962 (J. F. McAlpine); 2 males,
1 female, same locality, Aug. 7, 1961 (D. R. Oliver), ex snow
bunting nest; 2 males, Hazen Lake, John’s I., June 20, 1963 (B.
Hocking) and June 12, 1964 (R. E. Leech); 2 males, same locality,
June 20, 1963 (C. D. Sharplin). Melville I.: male, Bailey Point,
July 24, 1965 (J. E. H. Martin). Victoria I. 11 males, 8 females,
71° 177 N, 114° W, June 25-30, 1975 (G. and D. M. Wood).
Keewatin District: male, Repulse Bay, June 25, 1950 (J. E. H.
Martin); female, Chesterfield, July 18, 1950 (J. G. Chillcott).
Mackenzie District: 4 males, 2 females, 21 mi E Tuktoyaktuk,
July 17, 21, 1971 (D. M. Wood). Labrador: male, female,
Hebron, Aug. 5, 1974 (J. F. McAlpine), with puparia, nest of
savannah sparrow on rocky ledge in tundra. Quebec: male, Ft.
Chimo, Aug. 13, 1948 (H. N. Smith). Yukon Territory: 6 males,
7 females, Dempster Highway, mi 40, 51, 87, km 140.5, 155,
various dates June 17-Aug. 8, 1973, 1980, and 1982 (G. and D.
M. Wood); 2 females, Dempster Highway, km 155, June 29-July
3, 1980 (D. M. Wood, D. Lafontaine); male, 4 females, Dempster

P.tundrae 227

Highway, km 465, June 23-25, 1980 (D. Lafontaine). Greenland:
male, "Hekla-Havn," East Greenland, June 11, 1892 (H.
Deichmann) [Copenhagen Mus.]; male, Sondrestrom Air Base,
June 11, 1952 (W. J. Brown)[CNC]; 37 males, 31 females,
Pearyland, Nedre Midsommer S6 [Lake, N of head of
Independence Fjord], 82°14’ N, 33°20’ W, ex two snow bunting
nests, puparia July 5, 1966, adults July 10-20, 1966 (J. E. H.
Martin) [CNC].

Non-paratype Material: Two males, three females from the
Hazen Camp locality, a male from Pearyland, and a female from
Tuktoyaktuk, small or in poor condition.

Specimens Examined: 100 males, 100 females.

Distribution: Arctic Canada, Yukon Territory to southern
Greenland, Labrador and northern Quebec, north to Melville and
Ellesmere Islands and Pearyland in northern Greenland.

Basis of description of immature stages: Puparium (12
specimens): Hebron, Labrador, savannah sparrow, (2); Hazen
Camp, Ellesmere Island, N. W. T., snow bunting, (10).

Hosts: Snow bunting, savannah sparrow.

Ecology and Biology. Although records are few and only two
host species are known, this is clearly a species of the far north,
a tundra species, living in the nests of ground-nesting birds. The
records from snow bunting in Ellesmere Island and Pearyland in
northern Greenland are especially interesting, as this bird nests
"farther north than any other land bird" (AOU Checklist, 1957),
and our records are from the northernmost extent of its breeding
range.

Variation. Apparently the usual number of posterior bristles
on the fore tibia is two, but enough specimens have only one that
the variation will give some trouble to users of the keys. Of the
185 specimens that could be counted, 69 males and 50 females
have two posterior bristles on each fore tibia, and another 8
males and 30 females have two on one side or the other, but 6
males and 22 females have only one posterior bristle on each fore
tibia. One female (Dempster Highway) has one bristle on one
tibia but none at ail on the other.

Remarks. This species is very close to fallisi, but the
characteristic appearance of the frons and especially the
preocellar area is so consistent that we consider the two to be
distinct species. When puparia are compared, the hyperstigmatal
area in fallisi has stout, well spaced spines (Fig. 50), whereas in
tundrae these spines are relatively slender and appear to be closer
together (Fig. 64). Puparia of tundrae are most easily confused
with those of halli. They can be separated by the fact that the

228 _— Protocalliphora

posterior band, although reduced, is present on the anterior 5-6
segments of tundrae but absent on all segments of halli. In
addition, tundrae appears to be limited, distributionally, to the
arctic and tundra regions while the other species has only been
recorded in western North America.

The series of tundrae was compared with the type series (3
males, 7 females) of P. nuortevai Grunin from Finland, loaned
from the Zoological Museum of the University of Helsinki by
courtesy of B. Lindeberg. Unfortunately the sample of nuortevai
is small and its reared specimens are not in good condition. The
Nearctic and Palearctic specimens are very similar. The most
striking difference is the breadth of the parafacial in both sexes
of nuortevai, approximately twice the breadth of the 3rd antennal
segment, much greater than the ocellar span (2.00-2.14x in the
four measured females, 1.86 and 2.00x in the two measured
males), and in the four females slightly greater than breadth of
frons (0.54 and 0.555x). The surstyli are the same curved type.
The puparia are similar in gross appearance, although the few
specimens of nuortevai show stronger hypostigmatal spines.
Obviously tundrae and nuortevai are closely related, if indeed
they are not the same species with subspecific relationships.
Until a more adequate sample of nuortevai is available and
puparia are also studied, it seems best to recognize P. tundrae as
distinct. The far northern range of this species is remarkable.
The specimens from Ellesmere Island and Pearyland in northern
Greenland were reared from puparia found in snow bunting nests
at latitudes 81° 49’ N and 82° 14’ N, respectively.

Etymology. The specific name is in the genitive case, based
on the tundra habitat.

Calliphora splendida Macquart, species dubia

Calliphora splendida Macquart, 1846: 324 (1846: 196) [Galveston,
Tex. ].

Beginning with Harbeck (1907), this species was referred to
Protocalliphora, and was considered the senior synonym of P.
metallica (Townsend). The type of splendida, a female, has
apparently been lost. Townsend (1919), recognizing some
discrepancies of description, and Hall (1948), having regard to
the then unknown occurrence of Protocalliphora so far south,
both regarded splendida as_ unrecognizable. Today the
distribution is not quite as strong as argument, although the genus

Calliphora splendida 229

is still unknown from Texas (Cf. Map 1), and the likelihood of
its occurrence in coastal plain Texas is opposed by all we know
of the group. Furthermore, even if splendida were a
Protocalliphora, the association with metallica is by no means
certain. The principal color character, the cupreous apex of the
abdomen, is now known to be present in females of several
species of Protocalliphora, besides Neotropical flies of other
genera and families that might be involved. We prefer, with
Townsend and Hall, to leave Calliphora splendida as a species
dubia.

Bits
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ois

a Sa oN
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iy

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TR

Appendix 231

Appendix
Natural Hosts of Protocalliphora in North America

Common and scientific names are those adopted in the
"Check-List of North American Birds", 6th edition (1983),
published by the American Ornithologists’ Union. In a few
cases, well known former family names or common names for
species are cited in brackets immediately after the presently
adopted names. In addition to the natural hosts, the common
chicken and the budgerigar have been reported as hosts of
Protocalliphora (see under P. braueri), and P. sialia was reared
from a nest of caged ringed turtle-doves used in experimental
studies in Colorado, far from any feral established population of
the doves.

Birds are recorded by species only, not by subspecies, except
for the flickers, juncos, and yellow-rumped warbler, for which
relatively recent taxonomic changes have altered the long-used
classification and the common names. New host records are
starred (*), with the name of the collector(s) credited in brackets
at the end of the entry. The new records include those found in
the studies by Bennett and Whitworth, plus the hosts of reared
material identified by Sabrosky from various sources. References
are given for records overlooked in or published since the last
host list for North America (Hall 1948). Many of the published
hosts have also been recorded by Bennett or by Whitworth, who
reared Protocalliphora from at least 43 and 48 species,
respectively.

Blackbirds: Emberizidae, Icterinae [Icteridae]
Brewer’s, Euphagus cyanocephalus (Wagler) [Wood 1953, Bent
1958]
Red-winged, Agelaius phoeniceus (Linnaeus) [Miller 1909,
Judd 1954]
*Yellow-headed, Xanthocephalus xanthoce phalus (Bonaparte)
[Whitworth, C. S. Gold]

Bluebirds: Muscicapidae, Turdinae [Turdidae]
Eastern, Sialia sialis (Linnaeus)
*Mountain, Sialia currucoides (Bechstein) [G. J. Spencer,
Whitworth]
*Western, Sialia mexicana Swainson [Vasiliki Demas, C. S.
Gold]

232 ~~ Protocalliphora

Buntings: Emberizidae [Fringillidae, part], Cardinalinae (Indigo,
Lazuli) and Emberizinae (Snow)
Indigo, Passerina cyanea (Linnaeus)
*Lazuli, Passerina amoena (Say) [Whitworth]
*Snow, Plectrophenax nivalis (Linnaeus) [J. F. McAlpine, R.
B. Madge, J. E. H. Martin, D. R. Oliver, D. M. Wood]

Bushtits: Aegithalidae
Bushtit, Psaltriparus minimus (Townsend)

Cardinals: Emberizidae [Fringillidae, part], Cardinalinae
Northern, Cardinalis cardinalis (Linnaeus)

Catbirds: Mimidae
Gray, Dumetella carolinensis (Linnaeus)

Chats: Emberizidae, Parulinae [Parulidae]
Yellow-breasted, /cteria virens (Linnaeus)

Chickadees: Paridae

Black-capped, Parus atricapillus Linnaeus

*Carolina, Parus carolinensis Audubon [E. T. McKnight, W.
A. Tarpley]

Chestnut-backed, Parus rufescens Townsend [Gold &
Dahlsten 1984]

Mountain, Parus gambeli Ridgway [Storer 1929, Gold &
Dahlsten 1984]

Cowbirds: Emberizidae, Icterinae [Icteridae]
*Brown-headed, Molothrus ater (Boddaert) [V. E. Davis]

Creepers: Certhiidae
Brown, Certhia americana Bonaparte [Gold & Dahlsten 1984]

Crows: Corvidae
American [Common], Corvus brachyrhynchos Brehm

Cuckoos: Cuculidae
*Black-billed, Coccyzus erythropthalmus (Wilson) [Bennett]

Dippers: Cuinclidae
* American [Water Ouzel], Cinclus mexicanus Swainson [C. L.
Remington, Whitworth, J. A. Halstead]

Appendix 233

Doves: Columbidae
Mourning, Zenaida macroura (Linnaeus)

Eagles: Acciptridae
Bald, Haliaeetus leucocephalus (Linnaeus) [Bortolotti 1985, A.
H. Grewe, Jr., B. E. Cooper]
Golden, Aquila chrysaetos (Linnaeus) [Philip 1938, Hill and
Work 1947]

Falcons and Kestrels: Falconidae
American Kestrel [Sparrow Hawk], Falco sparverius Linnaeus
[Hill and Work 1947]
*Gyrfalcon, Falco rusticolus Linnaeus [K. G. Poole]
*Peregrine Falcon, Falco peregrinus Tunstall [J. A. Hagar]
Prairie Falcon, Falco mexicanus Schlegel [possible, White
1963]

Finches: Fringillidae
*Cassin’s, Carpodacus cassinii Baird [Whitworth]
House, Carpodacus mexicanus (Muller)
Purple, Carpodacus purpureus (Gmelin)
Rosy, Leucosticte arctoa (Pallas) [Black Rosy, L. atrata
Ridgway] [French 1959]

Flickers: Picidae
Northern [Common], Colaptes auratus (Linnaeus)
*Red-shafted, C. auratus cafer (Gmelin) [Whitworth]
Yellow-shafted, C. auratus auratus (Linnaeus)[Kenaga
1961]

Flycatchers: Tyrannidae

*Alder [Traill’s, in part], Empidonax alnorum Brewster
[Bennett]

*Ash-throated, Myiarchus cinerascens (Lawrence) [Vasiliki
Demas]

*Dusky, Empidonax oberholseri (Phillips) [C. S. Gold]

Great Crested, Myiarchus crinitus (Linnaeus)

*Western, Empidonax dif ficilis Baird [N. A. M. Verbeek, C.
S. Gold]

*Willow [Traill’s, in part], Empidonax traillii (Audubon) [C.
S. Gold, J. A. Halstead]

234 ~=‘Protocalliphora

Goldfinches: Fringillidae
American, Carduelis tristis (Linnaeus)
Lesser, Carduelis psaltria (Say)

Goshawks. See under Hawks

Grackles: Emberizidae, Icterinae [Icteridae]
*Common, Quiscalus quiscula (Linnaeus) [Bennett, D. M.
Davies, F. M. Stiner, Jr.]

Grosbeaks: Emberizidae, Cardinalinae, and Fringillidae (Pine
Grosbeak)
*Black-headed, Pheucticus melanocephalus (Swainson) [D. C.
Rentz]
Blue, Guiraca caerulea (Linnaeus) [Stabler 1959]
*Pine, Pinicola enucleator (Linnaeus) [Whitworth]

Harriers: See Hawks

Hawks: Acciptridae
Broad-winged, Buteo platypterus (Vieillot) [Meng 1954]
Cooper’s, Accipiter cooperii (Bonaparte)
*Ferruginous, Buteo regalis (Gray) [Whitworth]
Northern Goshawk, Accipiter gentilis (Linnaeus) [Meng 1954]
Northern Harrier [Marsh Hawk], Circus cyaneus (Linnaeus)
[F. & F. Hamerstrom 1954]
Red-shouldered, Buteo lineatus (Gmelin)
Red-tailed, Buteo jamaicensis (Gmelin)
Sparrow. See Falcons and Kestrels
Swainson’s, Buteo swainsoni Bonaparte

Jays: Corvidae
*Blue, Cyanocitta cristata (Linnaeus) [Bennett]
*Gray [Canada], Perisoreus canadensis (Linnaeus) [Bennett]
*Pinyon, Gymnorhinus cyanocephalus Wied [K. A. Walker]

Juncos: Emberizidae [Fringillidae, part], Emberizinae
Dark-eyed, Junco hyemalis (Linnaeus)
*Oregon, J. hyemalis oreganus (Townsend) [Whitworth]
Slate-colored, J. hyemalis hyemalis (Linnaeus)

Kestrels: See Falcons and Kestrels

Appendix 235

Kingbirds: Tyrannidae
Cassin’s, Tyrannus vociferans Swainson
*Eastern, 7 yrannus tyrannus (Linnaeus) [Bennett, Whitworth]
*Western [Arkansas], Tyrannus verticalis Say [Bennett,
Whitworth, G. J. Spencer]
Larks: Alaudidae
Horned, Eremophila alpestris (Linnaeus)

Magpies: Corvidae
Black-billed, Pica pica (Linnaeus)

Martins: Hirundinidae
Purple, Progne subis (Linnaeus)

Meadowlarks: Emberizidae, Icterinae [Icteridae]
Eastern, Sturnella magna (Linnaeus) [Miller 1909]
Western, Sturnella neglecta Audubon [Arnold 1919]

Mockingbirds: Mimidae
Northern, Mimus polyglottos (Linnaeus)

Nuthatches: Sittidae
Brown-headed, Sitta pusilla Latham [Norris 1958]
Pygmy, Sitta pygmaea Vigors [probable, Law 1929}
Red-breasted, Sitta canadensis Linnaeus [Gold & Dahlsten
1984]
*White-breasted, Sitta carolinensis Latham [D. L. Dahlsten;
Museum of Comparative Zoology, collector unknown]

Ospreys: Acciptridae, Pandioninae [Pandionidae]
*Osprey, Pandion haliaetus (Linnaeus) [F. and F.
Hamerstrom]

Ovenbirds: Emberizidae, Parulinae [Parulidae]
*Ovenbird, Seiurus aurocapillus (Linnaeus) [Bennett]

Owls: Strigidae
*Eastern Screech-, Otus asio (Linnaeus) [Bennett]
*Flammulated, Otus flammeolus (Kaup) [Whitworth]
Great Horned, Bubo virginianus (Gmelin) [Lee and Ryckman
1954, Tirrell 1978]
Long-eared, Asio otus (Linnaeus)
*Western Screech-, Otus kennicottii (Elliot) [Sabrosky]

236 —-Protocalliphora

Pewees: Tyrannidae
*Western Wood-, Contopus sordidulus Sclater [Whitworth]

Phoebes: Tyrannidae
*Black, Sayornis nigricans (Swainson) [H. B. Leech]
Eastern, Sayornis phoebe (Latham) [Brimley 1938]
Say’s, Sayornis saya (Bonaparte)

Pipits: Motacillidae
*Water, Anthus spinoletta (Linnaeus) [N. A. M. Verbeek]

Ravens: Corvidae
*Common, Corvus corax Linnaeus [Whitworth]

Redstarts: Emberizidae, Parulinae [Parulidae]
American, Setophaga ruticilla (Linnaeus) [Johnson 1930,
George and Mitchell 1948]

Robins: Muscicapidae, Turdinae [Turdidae]
American, Turdus migratorius Linnaeus

Shrikes: Laniidae
Loggerhead, Lanius ludovicianus Linnaeus

Siskins: Fringillidae
*Pine, Carduelis pinus (Wilson) [J. D. Tiner]

Solitaires: Muscicapidae, Turdinae [Turdidae]
Townsend’s, Myadestes townsendi (Audubon) [Munro 1949]

Sparrows: Emberizidae [Fringillidae, part], Emberizinae, and

Passeridae (House Sparrow)

American Tree, Spizella arborea (Wilson)

Chipping, Spizella passerina (Bechstein)

*Field, Spizella pusilla (Wilson) [V. Nolan, Jr.]

*Fox, Passerella iliaca (Merrem) [Whitworth]

*Grasshopper, Ammodramus savannarum (Gmelin) [D. F.
Owen]

House [English], Passer domesticus (Linnaeus)

Savannah, Passerculus sandwichensis (Gmelin) [Munro 1949,
Bedard and McNeil 1979]

Song, Melospiza melodia (Wilson)

Swamp, Melospiza georgiana (Latham)

*Vesper, Pooecetes gramineus (Gmelin) [Bennett]

Appendix 237

Sparrows: (continued)
White-crowned, Zonotrichia leucophrys (Forster)
White-throated, Zonotrichia albicollis (Gmelin)

Starlings: Sturnidae
European, Sturnus vulgaris Linnaeus

Swallows: Hirundinidae

Bank, Riparia riparia (Linnaeus)

Barn, Hirundo rustica Linnaeus

Cliff, Hirundo pyrrhonota (Vieillot)

*Northern Rough-winged, Stelgidopteryx serripennis
(Audubon) [S. ruficollis (Vieillot)] [Bennett]

Tree, Tachycineta bicolor (Vieillot)

*Violet-green, Tachycinetathalassina(Swainson)[Whitworth,
R. E. Leech, S. Marino and W. L. Jellison]

Tanagers: Emberizidae, Thraupinae [Thraupidae]
Scarlet, Piranga olivacea (Gmelin)

Thrashers: Mimidae
Brown, Toxostoma rufum (Linnaeus)

Thrushes: Muscicapidae, Turdinae [Turdidae]
Hermit, Catharus guttatus (Pallas) [George and Mitchell 1948]
Swainson’s[Olive-backed, Russet- backed], Catharus ustulatus
(Nuttall)
*Wood, Hylocichla mustelina (Gmelin) [Bennett, V. E. Davis,
C. Strehl, F. M. Stiner, Jr.]

Towhees: Emberizidae [Fringillidae, part], Emberizinae
Brown, Pipilo fuscus Swainson
Rufous-sided, Pipilo erythrophthalmus (Linnaeus)

Vireos: Vireonidae
*Red-eyed, Vireo olivaceus (Linnaeus) [Bennett]
Solitary [Blue-headed], Vireo solitarius (Wilson) [Johnson
1930]
*Warbling, Vireo gilvus (Vieillot) [Whitworth]

Warblers: Emberizidae, Parulinae [Parulidae]
Black-throated Blue, Dendroica caerulescens (Gmelin)
*Canada, Wilsonia canadensis (Linnaeus) [C. E. Hope]

238 Protocalliphora

Warblers: (continued)
Chestnut-sided, Dendroica pensylvanica (Linnaeus) [Johnson
1929a]
*Nashville, Vermivora ruficapilla (Wilson) [Bennett]
Prairie, Dendroica discolor (Vieillot) [Nolan 1955, 1959]
Wilson’s [Pileolated], Wilsonia pusilla (Wilson) [Rausch 1972]
Worm-eating, Helmitheros vermivorus (Gmelin)
Yellow, Dendroica petechia (Linnaeus)
*Yellow-rumped, Dendroica coronata (Linnaeus)
*Audubon’s, D. coronata auduboni (Townsend)
[Whitworth]
*Myrtle, D. coronata coronata (Linnaeus) [Bennett, C. E.
Hope]

Waterthrushes: Emberizidae, Parulinae [Parulidae]
*Louisiana, Seiurus motacilla (Vieillot) [A. Wetmore]
*Northern, Seiurus noveboracensis (Gmelin) [Bennett]

Waxwings: Bombycillidae
Cedar, Bombycilla cedrorum Vieillot

Woodpeckers: Picidae
*Downy, Picoides pubescens (Linnaeus) [Bennett]

Wrens: Troglodytidae
Bewick’s, Thryomanes bewickii (Audubon) [Gold & Dahisten
1981, 1984]
Carolina, Thryothorus ludovicianus (Latham)
House, Troglodytes aedon Vieillot
*Marsh [Long-billed Marsh], Cistothorus palustris (Wilson)
[Whitworth]

Yellowthroats: Emberizidae, Parulinae [Parulidae]
*Common, Geothlypis trichas (Linnaeus) [Bennett]

References 239

Annotated References

Annotations (by Sabrosky) in brackets give (1) the species as
recorded in the publication; (2) locality and host, unless these are
indicated in the title; and (3) the actual identity of the parasite
species as determined by rechecking original material, or the
probable identity if that can be deduced with reasonable
assurance from the published biology and distribution. The
species of blow flies referred to are all in Protocalliphora
(abbreviated as P.), unless otherwise stated. Only publications
that contain original records on Nearctic species are so annotated;
those that merely repeat earlier published records or information,
such as text books and some state lists, have not been included
unless they are cited for other reasons.

A few publications not specifically referred to in the text are
included because their host or locality records were used for lists,
tables, or maps.

Aldrich, John M.
1901. Synonymische Notiz. Wiener Entomologische Zeitung,
20: 68. [Avihospita Hendel 1901 = Protocalliphora Hough
1899}.
1905. A catalogue of North American Diptera. Smithsonian
Miscellaneous Collections, 46: 680 pp.
Allen, Robert W., and Margaret M. Nice
1952. A study of the breeding biology of the purple martin
(Progne subis). American Midland Naturalist, 47:
606-665. [Larvae of P. splendida, George Reserve near
Ann Arbor, Mich.; unverified].
American Ornithologists’ Union, Committee on Classification and
Nomenclature
1957. Check-List of North American birds (Sth edition).
America Ornithologists’ Union, pp. xiii + 691.
1983. Check-list of North American birds (6th edition).
American Ornithologists’ Union, Washington, D.C., pp.
XX1X + 877.
Arnold, W. W.
1919. Maggot infested birds. Auk, 36: 147-148. [P. azurea,
Colo., various hosts; = P. braueri, paratypes of hirudo].

240 ~— Protocalliphora

Banks, Nathan

1912. The structure of certain dipterous larvae with
particular reference to those in human foods. United
States Department of Agriculture, Bureau of Entomology,
Technical Series, 22: 44 pp. [P. chrysorrhoea, Penn. and
Mass., on bluebirds; = P. sialia.

Barbara Ann, Sister

1970. Five years’ experience with a bluebird population.
Maryland Birdlife, 26: 81-92. [Md.; P. sialia reared; det.
Sabrosky].

Bedard, Jean, and Jeremy N. McNeil

1979. Protocalliphora hirudo (Diptera: Calliphoridae)
infesting Savannah sparrow, Passerculus sandwichensis
(Aves: Fringillidae), in eastern Quebec. Canadian
Entomologist 111: 111-112. [det. G. F. Bennett and B.
Cooper].

Bennett, Gordon, F.

1957. Studies on the genus Protocalliphora (Diptera:
Calliphoridae). University of Toronto, Canada,
Unpublished Ph.D. Dissertation: 194 pp., 33 pls.

Bent, Arthur C.

1937. Life histories of North American birds of prey. Order
Falconiformes (Part 1). Bulletin of the United States
National Museum, 167: viii + 409, 102 pls.
[Protocalliphora larvae in ears of young red-shouldered
hawks, probably N.H.; undoubtedly P. avium].

1942. Life histories of North American flycatchers, lake:
swallows, and their allies. Bulletin of the United States
National Museum, 179: xi + 555, 70 pls. [p. 395, P.
splendida var. sialia, nest of tree swallow; p. 418, P.
splendida, Maine, bank swallow; p. 455, nest of barn
swallow].

1948. Life histories of North American nuthatches, wrens,
thrashers and their allies. Bulletin of the United States
National Museum, 195: xi + 475, 90 pls. [Protocalliphora
splendida sialia, citing Mason (1936)].

1958. Life histories of North American blackbirds, orioles,
tanagers, and allies. Bulletin of the United States
National Museum, 211: ix + 549, 37 pls. [Protocalliphora
larvae on nestlings of Brewer’s blackbird].

Bergtold, W. H.

1927. A house finch infected by fly larvae. Auk, 44:
106-107. [P. splendida, Colo., larvae in abscess;
undoubtedly P. braueri).

References 24]

Bezzi, Mario

1922: On the dipterous genera Passeromyia and
Ornithomusca, with notes and bibliography on the
non-pupiparous Myiodaria parasitic on_ birds.
Parasitology, 14: 29-46.

Bezzi, Mario and Paul Stein

1907. Band III. Cyclorrapha (part), 828 pp., in Becker,
Bezzi, Kertész, and Stein, Katalog der palaarktischen
Dipteren. 4 vols. Budapest.

Bohm, Robert T.

1978. Protocalliphora infestation in great horned owls.
Wilson Bulletin, 90: 297. [P. avium, det. Sabrosky, near
Foley, Minn., in ear cavities].

Bortolotti, Gary R.

1985. Frequency of Protocalliphora avium (Diptera:
Calliphoridae) infestations on bald eagles (Haliaeetus
leucocephalus). Canadian Journal of Zoology, 63:
165-168. [P. avium, det. Sabrosky, Sask., common in ear
cavities].

Boyd, A. W.

1935. A fly parasite on the swallow. British Birds, 28: 225.

1936. Report on the swallow enquiry, 1935. Ibid., 30:
98-116.

Boyes, J. W.

1961. Somatic chromosomes of higher Diptera. V.
Interspecific and intra-specific variation in the
Calliphoridae. Canadian Journal of Zoology, 39: 549-570,
1 plate. [6 spp. of Protocalliphora, Algonquin Park,
Ontario; "n. sp. near sialia" = shannoni].

Boyes, J. W., and Janny M. van Brink

1965. Chromosomes of Calyptrate Diptera. Canadian Journal
of Genetics and Cytology, 7: 537-550. [6 spp. of
Protocalliphora; "n. sp. near sialia" = shannoni].

Boyes, J. W., and G. E. Shewell

1975. Cytotaxonomy of Calliphoridae (Diptera). Genetica,
45: 435-488. [6 spp. of Protocalliphora; "n. sp. nr. sialia"
= shannoni].

Brimley, C. S.

1938. The insects of North Carolina. North Carolina
Department of Agriculture, Division of Entomology,
Raleigh, N.C., 560 pp. [P. avium, Raleigh, N.C., nest of
phoebe; = P. deceptor, paratypes].

242 ~~‘ Protocalliphora

Britton, W. E.

1920. Check-list of the insects of Connecticut. Bulletin of
the Connecticut State Geological and Natural History
Survey, 31: 397 pp. [P. azurea, Conn., apparently on
authority of C. W. Johnson].

Brower, A. E.

1966. Management of bird houses to reduce nesting loss to
Protocalliphora, a parasitic fly. Maine Field Naturalist,
22: 73-74. [No spp. mentioned; general on use of
pyrethrum powder in bird houses].

Burks, B. D.

1979, Family Pteromalidae, pp. 768-835, in Krombein et al.
(eds.), Catalog of Hymenoptera in America North of
Mexico, Vol. 1: 1198 pp. Washington, D.C.

Burtch, Verdi

1920. Maggots in the ears of nestling Cooper’s hawks
(Accipiter cooperi). Auk, 37: 293. [Larvae "possibly"
Compsomyia macellaria (Fabricius), Branchport, N.Y.;
presumably = P. avium].

Cais, Leszek

1965. [Protocalliphora hirudo Shannon and Dobroscky, 1924
(Diptera, Calliphoridae) in Poland] (In Polish). Fragmenta
Faunistica(Polska Akademia Nauk, Instytut Zoologiczny),
12: 183-191.

Chapman, B. R.

1973. The effects of nest ectoparasites on cliff swallow
populations. Texas Technological University,
Unpublished Ph.D. Dissertation: 70 pp. [Dissertation
Abstracts, B: 73-23753]. [No mention of Protocalliphora].

Chapman, L. B.

1935. Studies of a tree swallow colony. Bird-Banding,
6:45-57. [Protocalliphora larvae, Princeton, Mass.; not
checked].

Cole, Frank R., and A. L. Lovett

1921. An annotated list of the Diptera (flies) of Oregon.
Proceedings of the California Academy of Sciences, Series
4, 11: 197-344. [P. azurea, Corvallis, Oreg.; = P. braueri}.

Coutant, Albert F.

1915. The habits, life history, and structure of a
blood-sucking muscid larva (Protocalliphora azurea).
Journal of Parasitology, 1: 135-150. [Ithaca, N.Y., crow;
= P. avium].

References 243

Crocoll, Scott, and James W. Parker

1981. Protocalliphora infestation in broad-winged hawks.
Wilson Bulletin, 93: 110. [Chautauqua Co., N.Y.; P.
avium].

Crosskey, R. W.

1981. Simuliid taxonomy--the contemporary scene. pp.
3-18, in Laird, M. (ed.). Blackflies. Academic Press,
New York, pp. xiv + 399.

Dobroscky, Irene D.

1925. External parasites of birds and the fauna of birds’
nests. Biological Bulletin, 48: 274-281 [P. avium, from
crows, and P. splendida, N.Y.; avium probably correct,
latter probably metallica}.

Dodge, Harold R., and John M. Seago

1954. Sarcophagidae and other Diptera taken by trap and net
on Georgia mountain summits in 1952. Ecology, 35:
50-59. [Protocalliphora, 2 spp., not named; = P. sialia
and P. braueri].

Dufour, L.

1845. Histoire des métamorphoses de la Lucilia dispar.
Annales de la Société Entomologique de France, Séries 2,
3: 205-214. [Earliest biological study of bird blow flies,
now considered to be P. azurea (Fallén)].

Emden, F. I. van

1954. Diptera Cyclorrhapha, Calyptrata (I), Section (a)
Tachinidae and Calliphoridae. pp. 133, Jn Handbooks for
the Identification of British Insects, vol. X, Part 4a.

Engel, Erich O.

1920. Dipteren, die nicht Pupiparen sind, als Vogelparasiten.
Zeitschrift fiir Wissenschaftliche Insektenbiologie, 15:
249-258, 9 figs.

Eshuis-van der Voet, C. W.

1972. Parasitism by Protocalliphora azurea (Fall.).
Verhandelingen der Koninklijke Nederlandsche
Akademie van Wetenschappen, Afdeeling Natuurkunde,
Tweede Reeks (Series 2), 61(3): 73-74.

1975. Parasitism by Protocalliphora spp. Ibid. 66: 81-84.

Eshuis-van der Voet, C. W., and E. Houwink
1976. Parasitism by Protocalliphora spp. Ibid. 67: 116.
Eshuis-van der Voet, C. W., and H. N. Kluyver

1971. Parasitism by Protocalliphora azurea Fall. Ibid. 60(2):

50-51.

244 ~~ Protocalliphora

Eshuis-van der Voet, C. W., and R. H. de Reede

1974. The effect of parasitism by Protocalliphora. Ibid. 63:

77-79.
Fallén, C. F.

1817. Beskrifning 6fver de i Sverige funna Fluge Arter, som
kunna foras till slagtet Musca. Foérsta Afdelningen.
Kongliga Vetenskaps- Academiens MHandlingar
(Stockholm), [ser. 3], 1816: 226-254. [Musca azurea, Nn.
sp., p. 245].

1821. Monographia Muscidum Sveciae (part). pp. 41-48.
Lund, Sweden.

Fan, C. T.

1965. [Key to the common synanthropic flies of China] (In
Chinese). Academy of Science, Peking, pp. xv + 330, 40
pls.

French, Norman R.

1959. Life history of the Black Rosy Finch. Auk, 76:
161-180. [NW Wyoming; Protocalliphora new species =
P. spatulata n. sp.].

Gahan, A. B.

1933. Description of a chalcidoid parasite of Protocalliphora
(Hymenop.). Canadian Entomologist, 65: 31-33.
[Morodora armata new genus and species, from pupae of
P. avium, Ravalli County, Mont.; = P. asiovora].

George, J. L., and R. T. Mitchell

1948. Notes on two species of Calliphoridae (Diptera)
parasitizing nestling birds. Auk, 65: 549-552. [Apaulina
metallica and A. hirudo, Lake Clear Junction, N.Y.;
former is chiefly P. metallica, partly P. bicolor; P. hirudo
(now braueri) is correct].

Gibson, Arthur

1917. The entomological record, 1916. Annual Report of the
Entomological Society of Ontario, (1916) 47: 137-171. [P.
azurea, Ottawa, Ont.; = P. metallica}.

1919. The entomological record, 1918. Annual Report of the
Entomological Society of Ontario, (1918) 49: 97-123.
[Phormia azurea, Outremont, Que.; not checked].

Gill, Gordon D.

1956. Filth flies of central Alaska. Journal of Economic
Entomology, (1955) 48: 648-653 [P. avium, det. M. T.
James; not checked].

References 245

Goddard, Jerome, and Paul K. Lago

1983. An annotated list of the Calliphoridae (Diptera) of
Mississippi. Journal of the Georgia Entomological
Society, 18(4): 481-484 [P. n. sp. = P. deceptor).

Gold, C. S., and D. L. Dahlsten

1981. A new host record for Tachinaephagus zealandicus
[Hym.: Encyrtidae]. Entomophaga, 26: 459-460 [Host: P.
Nn. Sp., puparia only].

1984. Effects of parasitic flies (Protocalliphora spp.) on
nestlings of mountain and chestnut-backed chickadees.
(1983) Wilson Bulletin, 95(4): 560-572, 4 tables [P. n. sp.
no. | = P. parorum].

Gregor, Frantisek, and Dalibor Povolny

1959. Kritischer Beitrag zur Kenntnis der Tribus Phormiini
(Diptera, Calliphoridae). Casopis Ceskoslovenské
Spolecnosti Entomologické, 56: 26-51, 5 plates.

Grimshaw, Percy H.

1901. Diptera. Jn Fauna Hawaiiensis (David Sharp, editor),
3 (part i):1-77, 3 pls. [Calliphora azurea, Kona, Hawaii;
= Phormia regina (Meigen)].

Grunin, K. J.

1966. [New and little-known Calliphoridae (Diptera), mainly
blood-sucking or subcutaneous parasites of birds] (In
Russian). Entomologicheskoe Obozrenie, 45: 897-903
[English translation, Entomological Review 45: 503-506].

1970a. 108. Fam. Calliphoridae. pp. 607-624, in [Flies, Fleas]
(Stackelberg and Nartshuk, editors), volume 5, part 2, of
Bei-Bienko, G. J. (editor), [Identification of the insects
of the European part of the USSR] (In Russian).
Zoological Institute of the Academy of Sciences,
Leningrad, 943 pp.

1970b. [New species of Calliphoridae (Diptera) for the fauna
of the USSR] (In Russian). Entomologischeskoe
Obozrenie, 49: 471-483.

197la. Ergebnisse der zoologischen Forschungen von Dr. Z.
Kaszab in der Mongolei. 277. Calliphoridae (Diptera) II.
Acta Zoologica Academiae Scientiarum Hungaricae, 17:
255-259.

1971b. [Phormiata Grunin, gen.n.--The sixth genus of the
tribe Phormiini (Diptera, Calliphoridae)] (In Russian).
Entomologicheskoe Obozrenie, 50: 444-445.

246 —_ Protocalliphora

1972. Beschreibung einer ornithoparasitischen Fliege,
Protocalliphora nuortevai Grunin, sp. n. (Dipt.,
Calliphoridae) aus Nord-Finnland. Annales Entomologici
Fennici, 38: 156-158.

1975. [Gasterophilidae, Calliphoridae, Oestridae,
Hypodermatidae (Diptera) of the Soviet-Mongolian
expedition, 1969-1971] (In Russian). [Insects of
Mongolia], 3: 620-627. Zoological Institute, Academy of
Sciences of the USSR.

Grunin, K. J., P. Nuorteva, and P. Rajala

1969. Trypocalliphora lindneri Peus (Dipt., Calliphoridae) as
a subcutaneous parasite of the wheatear in northern
Finland. Annales Entomologici Fennici, 35: 56-57.

Guberlet, J. E., and H. H. Hotson

1940. A fly maggot attacking young birds, with observations
of its life history. Murrelet, 21: 65-68. [P. hesperia,
Seattle, Wash., subcutaneous infestation of house sparrow;
= P, braueri].

Hagar, Joseph A.

1969. History of the Massachusetts peregrine falcon
population, 1935-57. pp. 123-131, in Hickey, J. J.
(editor), Peregrine falcon populations: their biology and
decline. University of Wisconsin Press, Madison, Wis.,
pp. xxii + 596. [Protocalliphora larvae in ears;
undoubtedly P. avium].

Hakanen, R., K. J. Grunin, and P. Nuorteva

1974. Larvae of Trypocalliphora lindneri Peus (Dipt.,
Calliphoridae) as subcutaneous pathogens on nestlings of
the meadow pipit and common redpoll in the subarctic.
Annales Entomologici Fennici, 40: 15-18.

Hall, David G.

1948. The blowflies of North America. Thomas Say
Foundation (Entomological Society of America),
Lafayette, Ind., 477 pp., 51 plates. [10 spp. of Apaulina,
1.e., Protocalliphora; see records under individual species;
exact date of publication noted in Sabrosky, 1950].

1965. Family Calliphoridae [except Melanomyini]. pp.
922-933, in Stone, Alan, et al., A catalog of the Diptera
of America north of Mexico. United States Department
of Agriculture, Agriculture Handbook 276: pp. iv + 1696.

Hamerstrom, Frances, and Frederick Hamerstrom

1954. Myiasis of the ears of hawks. Falconry News and
Notes 1 (3): 4-8 [Protocalliphora, lowa, Wis., Mich.;
undoubtedly P. avium].

References 247

Harbeck, H. S.

1907. [Seven species of Diptera new to the New Jersey state
list of insects]. Entomological News, 18: 216. [P.
splendida, Ashland, N.J.; not checked].

Hardy, D. Elmo

1981. Diptera Cyclorrhapha IV. pp. vii + 491. Jn Insects of

Hawaii, vol. 14. University Press of Hawaii, Honolulu.
Hearle, Eric

1938. Insects and allied parasites injurious to livestock and
poultry in Canada. Canada Department of Agriculture,
Farmers’ Bulletin, 53: 108 pp. [P. hirudo cuprea, eastern
Canada, in tumorlike swellings on swallows, P. sp. on
baby chicks, and P. avium, eastern Canada, on swallows;
none checked: first two undoubtedly braueri, but avium
doubtful on swallows].

Heinz, H. J.

1954. Weitere Vogelblutfliegen (Gattung Protocalliphora,
Dipt.) in Deutschland und Osterreich. Die Vogelwarte, 17:
39-42, 3 figs. [P. hirudo (now braueri) in Austria,
Steiermark].

Hendel, Friedrich

1901. Beitrag zur Kenntnis der Calliphorinen (Dipt.). Wiener

Entomologische Zeitung, 20: 28-33.
Hennig, Willi _

1939. Uber Namen und Artenzahl der deutschen
"Vogelblutfliegen." Arbeiten ttber Physiologische und
Angewandte Entomologie aus Berlin-Dahlem, 6: 359-364.

1952. Die Larvenformen der Dipteren, Part 3.
Akademie-Verlag: Berlin, pp. vii + 628.

Henshaw, H. W.

1908. A parasitic fly injurious to our native birds. Auk, 25:
87-88. [P. chrysorrhoea, Wellesley Hills, Mass., on
bluebirds; = P. sialia, part of type series].

Hickey, Joseph J.

1942. Eastern population of the duck hawk. Auk, 59:
176-204. [Protocalliphora larvae, southern N.Y., on
peregrine falcon; probably P. avium].

1969(editor). Peregrine falcon populations: Their biology and
decline. University of Wisconsin Press, Madison, Wis.,
pp. xii + 596, 60 plates. [Plate 43: Protocalliphora larvae,
Vt., in auditory meatuses of young peregrine falcons;
undoubtedly P. avium].

248 Protocalliphora

Hicks, Ellis A.

[1959]. Check-list and bibliography on the occurrence of
insects in birds’ nests. Iowa State College Press, Ames,
Iowa, 681 pp.

1962. Ibid. Supplement I. Iowa State Journal of Science,
36(3): 233-348.

1971. Ibid. Supplement II. Iowa State Journal of Science,
46(2): 123-338.

Highhouse, W. L.

1963. Operation bluebird (1962) in Warren County, Penn.
The Kingbird, 13(2): 24. [Apaulina larvae; use of flea
powder in nests].

Hill, H. M.

1948. Raptorial hosts of Protocalliphora. Condor, 50: 131.

[Corrections to Hill and Work (1947)].
Hill, H. M., and T. H. Work

1947. Protocalliphora \arvae infesting nestling birds of prey.
Condor, 49: 74-75. [Protocalliphora larvae, Alameda Co.,
Calif., sparrow hawk and golden eagle; probably P.
asiovora].

Horning, Donald S., Jr., and William F. Barr

1970. Insects of Craters of the Moon National Monument
Idaho. Miscellaneous Series of the University of Idaho
College of Agriculture, no. 8: 118 pp. [Protocalliphora n.
spp. A., B, and sp. = P. seminuda, P. beameri, and P.
brunneisquama, respectively].

Hough, Garry de N.

1899a. Some North American genera of the dipterous group,
Calliphorinae Girschner. Entomological News, 10: 62-66
(March).

1899b. Synopsis of the Calliphorinae of the United States.
Zoological Bulletin, 2: 283-290 (Sept.).

ICZN (International Code of Zoological Nomenclature)

1961 (1st edition), 1985 (third edition). International Trust

for Zoological Nomenclature, London, England.
Ishijima, Hideo

1967. Revision of the third stage larvae of synanthropic flies
of Japan (Diptera: Anthomyiidae, Muscidae,
Calliphoridae and Sarcophagidae). Japanese Journal of
Sanitary Zoology, 18: 47-100, 12 plates. [P. azurea. brief
description of larva, figures of cephalopharyngeal
sclerites and posterior spiracles].

References 249

Itamies, Juhani, and Mikko Ojanen

1975. Trypocalliphora lindneri Peus. (Dipt. Calliphoridae)
infesting a great tit nestling (Parus major L.). Annales
Entomologici Fennici, 41: 42.

James, Maurice T.

1955. The blowflies of California. Bulletin of the California
Insect Survey, 4 (1): 34 pp. [7 species of Protocalliphora,
with Calif. records for 5 (see under individual species);
"hesperia" is a mixture of three species].

1969. A study in the origin of parasitism. Bulletin of the
Entomological Society of America, 15: 251-253.

Jellison, William L.

1949. Biological studies on the faunae of nests of birds and
rodents in relation to disease of animals and man.
University of Minnesota, Summaries of Ph.D. Theses, 3:
53-57. [Thesis of 1940, Protocalliphora on crow and
magpie fledglings; = P. asiovora].

Jellison, William L., and Cornelius B. Philip

1933. Faunae of nests of the magpie and crow in western
Montana. Canadian Entomologist, 65: 26-31 [P. avium;
= P. asiovoral].

Johannsen, O. A.

1928. Diptera, pp. 687-868, in Leonard, M. D. (editor), A
list of the insects of New York. Memoir of the Cornell
University Agricultural Experiment Station, 101: 1121 pp.
[Protocalliphora, p. 827, N.Y. records for P. avium, P.
splendida form splendida, and splendida form sialis (sic);
P. avium verified, form splendida not checked, of form
sialia the males of the McLean, Caroline, and Fall Creek
records are P. metallica, the Farmingdale specimen not
seen].

Johnson, Charles W.

1910. Order Diptera, pp. 703-814, in Smith, John B., The
insects of New Jersey. Annual Report of the New Jersey
State Museum 1909: 15-888 [P. splendida, Ashland, N.J.,
July 16; not checked].

1925a. Fauna of New England. 15. List of the Diptera or
two-winged flies. Occasional Papers of the Boston
Society of Natural History, 7: 326 pp. [P. splendida = part
P. metallica, part P. bicolor; P. hirudo cuprea = P.
braueri; P. splendida sialis [sic] = part P. sialia, P.
shannoni, P. metallica}.

250 —- Protocalliphora

1925b. Insects that infest birds. Bulletin of the Northeastern
Bird-Banding Association, 1: 51-53. [P. hirudo cuprea,
Middleboro, Mass., in neck of sparrow; = P. braueri].

1927a. The infestation of bluebirds’ nests by Protocalliphora.
Bulletin of the Northeastern Bird-Banding Association,
3: 1-3 [P. splendida form sialia, Rock and Needham,
Mass.; = P. sialia].

1927b. Dipterological notes. Psyche, 34: 33-35. [Same
records as in 1927a].

1927c. Further studies of Protocalliphora infesting nestling
birds. Bulletin of the Northeastern Bird-Banding
Association, 3: 77-82. [P. splendida splendida, Ashland,
N. H., black-throated blue warbler, and P. splendida
form sialia, various localities and hosts; former = P.
bicolor, latter = P. sialia].

1927d. The insect fauna. Biological survey of the Mount
Desert region, Part 1: 247 pp. [P. splendida form sialia,
Southwest Harbor, Maine; not checked; same record
repeated in enlarged editions of 1938 and 1946].

Johnson, Charles W.

1929a. Some additional notes on Protocalliphora. Bulletin of
the Northeastern Bird-Banding Association, 5: 29-30. [P.
splendida splendida, N.H., chestnut-sided warbler and
robin = P. bicolor and P. shannoni, respectively; P.
splendida var. sialia, Mass., bluebird and barn swallow,
verified as sialia; P. hirudo var. cuprea, Mass. = braueri,
repetition of Johnson, 1925b].

1929b. The injury to nestling birds by the larvae of
Protocalliphora. Annals of the Entomological Society of
America, 22: 131-135. [Review of the 1925-1928
seasons].

1930. Notes on Protocalliphora during the summer of 1930.
Bird-Banding, 1: 169-173. [P. sialia, Mass., nests of
bluebird and tree swallow = P. sialia; P. splendida
splendida, N.H., nests of redstart and blue-headed vireo
= P. bicolor].

1931. Nestling birds destroyed by the larvae of
Protocalliphora. Bulletin of the Boston Society of Natural
History, 59: 21-24. [Brief review of previous records].

1932. Notes on Protocalliphora during the summer of 1931.
Bird-Banding, 3: 26-29 [P. sialia, Mass., Vt., Ohio = P.
sialia; P. splendida, Mass. = P. metallica].

References 251

Judd, W. W.

1944. Dipterous larvae parasitic on nestlings of the song
sparrow. Canadian Field Naturalist, (1943) 57: 146. [P.
splendida, Que.; = P. metallica].

1954. Insects collected from birds’ nests at London, Ontario.
Canadian Field Naturalist, 68: 122-123. [P. sp. near
sialia, red-winged blackbird; = shannoni].

1957. Diptera (Calliphoridae, Heleidae, and Chloropidae)
collected from birds’ nests at London, Ontario.
Entomological News, 68: 32-34. [P. sp. (sialia?), robin
and brown thrasher; = P. shannoni].

Kamal, Adel S.

1958. Comparative study of thirteen species of
sarcosaprophagous Calliphoridae and Sarcophagidae
(Diptera). I. Bionomics. Annals of the Entomological
Society of America, 51: 261-271. [P. avium, Moscow,
Idaho, on magpies; undoubtedly P. asiovora].

Kano, Rokuro, and Satoshi Shinonaga

1966. Notes on flies of medical importance in Japan, Part
XXV. Descriptions of two species belonging to the genus
Protocalliphora(Diptera: Calliphoridae). Japanese Journal
of Sanitary Zoology, 17: 164-168.

Kenaga, Eugene E.

1961. Some insect parasites associated with the eastern
bluebird in Michigan. Bird-Banding, 32: 91-94. [P.
sialia, det. Sabrosky; also Protocalliphora larvae recorded
on other birds].

Kluyver, H. N., and C. W. Eshuis-van der Voet

1970. Parasitism by Protocalliphora sp. Verhandelingen der
Koninklijke Nederlandsche Akademie van
Wetenschappen, Afdeeling Natuurkunde, Tweede Reeks
(Series 2), 59(3): 48.

Knowlton, George F.

1936. Some Utah flesh flies. Proceedings of the Utah
Academy of Sciences, Arts and Letters, 13: 237-239. [P.
splendida, Hooper and Salt Lake City, Utah; Hooper
specimen not located but possibly cuprina, Salt Lake City
specimen = Phormia regina (Meigen)].

Krug, Howard H.

1941. Bluebird banding at Chesley, Ontario. Bird-Banding,
12: 23-26. [Maggots, probably Protocalliphora, in nests of
bluebirds and tree swallows].

252 ~~ Protocalliphora

Laskey, Amelia R.

1948. Some nesting data on the Carolina wren at Nashville,
Tennessee. Bird-Banding, 19: 101-121. [Pupa cases of
Protocalliphora}.

Law, J. Eugene

1929. Down-tree progress of Sitta pygmaea. Condor, 31:
45-51. [Flies, presumably Protocalliphora, entering nests,
Calif.].

Lee, Robert D., and Raymond E. Ryckman

1954. Coleoptera and Diptera reared from owl nests.
Bulletin of the Brooklyn Entomological Society, 49:
23-24. [Protocalliphora sp., Flagstaff, Ariz., nest of great
horned owl; = P. asiovora].

Lehrer, Andy Z.

1970. Considérations phylogénétiques et taxonomiques sur la
famille Calliphoridae (Diptera). Annotationes Zoologicae
et Botanicae, no. 61: 1-51, 44 figs.

Lincoln, F. C.

1931. Protocalliphora infestation of barn and bank swallows.
Bird-Banding, 2: 33-34. [P. splendida larvae, Fairdale, N.
Dak., barn swallows; very small 3rd-instar larvae,
probably sialia but possibly hirundo].

Lindner, Erwin

1957. Vogelblutfliegen (Protocalliphora, Calliphorinae,
Diptera): Ein Beitrag zur Kenntnis ihrer Systematik und
Biologie. Die Vogelwarte, 19: 84-90. .

Linsdale, J. M.

1937. The natural history of magpies. Cooper Ornithological

Club, Pacific Coast Avifauna, 25: 234 pp.
Lloyd, Gary D., and Cornelius B. Philip

1966. The "wingless" fly, Carnus hemapterus Nitzsch
(Milichiidae), on hawk fledglings in northern Utah.
Journal of Parasitology, 52: 414. [Protocalliphora larvae,
heavy infestations in ears of nestling red-tailed hawks,
Summit Co., Utah; undoubtedly P. asiovora].

Lloyd, Hoyes

1922. Larvae of Phormia chrysorrhea [sic] Macq. found upon
nestling bluebirds. Canadian Field Naturalist, 36: 116.
[Near Hull, Que.; = P. sialia}.

1927. Identification of flies found infesting bluebirds.
Canadian Field Naturalist, 41: 111. [Correction of 1922
identification to Protocalliphora splendida; = sialia].

References 253

Lundbeck, William

1895. QOstgrgnlandske Insekter. Fortegnelse over de
indsamlede _Insekter. Meddelelser om Grgnland
(Copenhagen), 19: 105-120. [Calliphora azurea (Fallén),
Hekla Havn, East Greenland; = P. tundrae. Same record
repeated by Lundbeck, 1900, on the Diptera of
Greenland, and by Henriksen and Lundbeck, 1917, on the
insects of Greenland].

Macquart, Jean

1846. Diptéres exotiques nouveaux ou peu connus. [I°"]
Supplément. Mémoires de la Société Royale des Sciences,
de l’Agriculture et des Arts de Lille, 1844: 133-364, 20
plates (Also separate publication, 1846, Paris, pp. 5-238,
20 plates). [Calliphora splendida, n. sp. Texas; = species
dubia].

Marshall, A. G.

1981. The ecology of ectoparasitic insects. New York,

Academic Press. Pp. xvi + 459.
Mason, Edwin A.

1936. Parasitism of birds’ nests by Protocalliphora at Groton,
Massachusetts. Bird-Banding, 7: 112-121. [Larvae of
Protocalliphora spp., various hosts].

1944. Parasitism by Protocalliphora and management of
cavity-nesting birds. Journal of Wildlife Management, 8:
232-247. [P. splendida, Groton, Mass., various hosts; =
mixture of sialia, shannoni, and metallica].

McAlpine, J. F., et al.

1981. Manual of Nearctic Diptera. Volume 1. Research
Branch, Agriculture Canada, Monograph 27: 674 pp.

1987. Ibid. Volume 2. Research Branch, Agriculture
Canada, Monograph 28: 1332 pp. (Calliphoridae by G. E.
Shewell, pp. 1133-1145).

McAtee, W. L.

1927. Notes on the insect inhabitants of bird houses.
Proceedings of the Entomological Society of Washington,
29: 87-90. ["Sarcophagid puparia" found in many nests,
Bell, Md.; = chiefly Protocalliphora, according to McAtee
1929].

1929. Further notes on insect inhabitants of bird houses.
Proceedings of the Entomological Society of Washington,
31: 105-111. [P. splendida, Bell, Md., 5 hosts; = P. sialia
in nests of starling, house sparrow, and purple martin,
other hosts not checked; P. sp., probably new, from nest
of house wren, = P. deceptor, n. sp.].

254 _ Protocalliphora

1940. An experiment in song bird management. Auk, 57:
333-348. [Protocalliphora larvae, Glenn Dale, Md., in
purple martin house].

Meigen, Johann W.

1824. Systematische Beschreibung der bekannten
europdischen zweifligeligen Insekten. Vol. 4: xii + 428
pp, pls. 34-41. Hamm.

1826. Systematische Beschreibung der bekannten
europaischen zweifligeligen Insekten. Vol. 5: xii + 412
pp., pls. 42-54. Hamm.

Meng, Heinz

1954. The bird nest screw-worm fly (Apaulina avium).
Falconry News and Notes, 1(5): 21-22. [Myiasis by A.
avium in ears of hawks, N.Y.; = undoubtedly P. avium].

Miller, Charles W.

1909. The occurrence of the larvae of a parasitic fly,
Protocalliphora chrysorrhoea, Meigen, in bird nests in the
vicinity of Shawnee, Pa. Bulletin of the Worthington
Society for the Study of Bird Life, 2: 8 pp. [Attacking
various birds; reared adults from bluebird nests = P.
sialia].

Miller, Robert C.

1929. Subcutaneous parasitism of a young warbler by fly
larvae. Murrelet, 10(1): 13-14. [P. splendida var. hesperia
infesting yellow warbler, Seattle, Wash., = P. braueri].

Munro, J. A.

1949. The birds and mammals of the Vanderhoof region,
British Columbia. American Midland Naturalist, 41:
1-138. [P. hirudo (now braueri) abundant; adults reared
from song sarrow = braueri, but larvae from some dead
fledglings proved to be Phaenicia sericata (Meigen) and
Phormia regina (Meigen)].

Neff, Johnson A.

1945. Maggot infestation of nestling mourning doves.
Condor, 47: 73-76. [P. metallica and an. sp., Calif.; =
cuprina and asiovora (synonym, basingeri Hall),
respectively].

Nielsen, J. C.

1907. The insects of East-Greenland. Meddelelser om
Grenland, 29: 365-409. [Calliphora azurea (Fallén),
record of Lundbeck (1895) repeated, with addition of
exact date, 11 June 92; = P. tundrae].

References 255

Nielsen, Peder, Oscar Ringdahl, and S. L. Tuxen

1954. Diptera 1 (exclusive of Ceratopogonidae and
Chironomidae). 189 pp., in The Zoology of Iceland, Vol.
III, Part 48a. Copenhagen and Reykjavik.

Nolan, Val, Jr.

1955. Invertebrate nest associates of the prairie warbler.
Auk, 72: 55-61. [P. metallica, Bloomington, Ind.; det.
Sabrosky].

1959. Additional invertebrate nest associates of the prairie
warbler. Auk, 76: 352-357. [P. metallica and a N. sp.,
Bloomington, Ind.; = P. metallica and P. deceptor].

1978. The ecology and behavior of the prairie warbler,
Dendroica discolor. Ornithological Monographs
(American Ornithologists’ Union), 26: xxii + 595,
frontispiece (color), 42 figs. [Updating of records of
species recorded in Nolan (1955, 1959)].

Norris, Robert A.

1958. Comparative biosystematics and life history of the
nuthatches Sitta pygmaea and Sitta pusilla. University
of California Publications in Zoology, 56: 119-300. [P.
"near sialia" (det. W. W. Wirth), one puparium, Irwinville,
Ga., brown-headed nuthatch nest; not checked].

Nuorteva, Pekka

1960. Protocalliphora chrysorrhoea (Meig.) karpasen toukat
tormapaaskyn loisina (Protocalliphora chrysorrhoea
(Meig.) as a parasite of Riparia riparia (L.) in Finland).
Ornis Fennica, 37: 122-124.

Nuorteva, Pekka, and Unto Jarvinen

1961. The insect fauna of the nests of the sand martin
(Riparia riparia L.) in Finland. Annales Entomologici
Fennicae, 27: 197-204. [Protocalliphora chrysorrhoea
(Meigen)].

Oliver, Don R.

1963. Entomological studies in the Lake Hazen area,
Ellesmere Island, including lists of species of Arachnida,
Collembola, and Insecta. Arctic (Journal of the Arctic
Institute of North America), 16: 175-180. [P. sp.; = P.
tundrae}.

Owen, D. F.

1954. Protocalliphora in birds’ nests. British Birds, 47:
236-243.

Owen, D. F., and J. S. Ash

1955. Additional records of Protocalliphora (Diptera) in
birds’ nests. British Birds, 48: 225-229.

256 —_ Protocalliphora

Peck, O.

1963. A catalogue of the Nearctic Chalcidoidea (Insecta:
Hymenoptera). Canadian Entomologist-Supplement,
1963: 1092 pp.

Petch, C. E., and J. B. Maltais

1932. [Revision of] A preliminary list of the insects of the
province of Quebec, Part II, Diptera, by A. F. Winn and
G. Beaulieu. Annual Report of the Quebec Society for
the Protection of Plants, 24, Supplement: 100 pp. [P.
splendida, Aylmer, Que. = P. metallica; P. azurea,
Montreal, not checked].

Peterson, Roger Tory

1947. A field guide to the birds (ed. 2). Houghton Mifflin

Co., Boston, pp. xxvi + 290.
Peus, Fritz

1960. Zur Kenntnis der ornithoparasitischen Phormiinen
(Diptera, Calliphoridae). Deutsche Entomologische
Zeitschrift, Neue Folge, 7: 193-235.

Philip, Cornelius B.

1938. A parasitological reconnaissance in Alaska with

particular reference to varying hares. II. Parasitological

data. Journal of Parasitology, 24: 483-488.
[Protocalliphora larvae, Rapids, Alaska, golden eagle; =
P. braueri].

Plath, O. E.

1919a. Parasitism of nestling birds by fly larvae. Condor, 21:
30-38 (January). [P. azurea, San Francisco Bay area,
various hosts; = hesperia complex].

1919b. A muscid larva of the San Francisco Bay region
which sucks the blood of nestling birds. University of
California Publications in Zoology, 19: 191-200
(February). [P. azurea, same material as in Plath, 1919a].

1919c. The prevalence of Phormia azurea Fallen (larva
parasitic on nestling birds) in the Puget Sound region and
data on two undescribed flies of similar habit. Annals of
the Entomological Society of America, 12: 373-378
(December)(Phormia metallica Townsend described on
pp. 379-380. See Townsend 1919). [Phormia azurea and
Phormia metallica; part of azurea = paratypes of
Protocalliphorahesperiaand hes perioides,n.sp.; metallica
from robin, Puget Sound, formed type series of
Protocalliphora hirudo var. cuprea, which equals braueri,
plus several females of cuprina].

References 257

Pletsch, Don D.

1948. Parasitic dipterous larvae from the nasal cavity of a
nestling magpie. Auk, 65: 296-297. [P. avium, Bozeman,
Mont.; = P. asiovora].

Pont, Adrian C.

1976. Calliphoridae, pp. 107-108, in Kloet and Hincks, A
Check List of British Insects, 2nd edition, Volume XI,
Part 5: Diptera and Siphonaptera.

Poole, K.G., and R.G. Bromley

1988. Natural history of the gyrfalcon in the Central
Canadian Arctic. Arctic, 41: 31-38. [P. avium, Kilgavik,
Mackenzie District, Northwest Territories, larvae].

Rausch, Robert L.

1972. Cutaneous myiasis in a bird by the _ larval
Protocalliphora h. hirudo Shannon and Dobroscky, 1924
(Diptera: Calliphoridae). Aquilo, Series Zoologica, 13:
1-4, [P. hirudo, Alaska, on Wilson’s warbler; = braueri].

Ringdahl, Oscar

1937. Bidrag till kinnedomen om de svenska tachinidernas
utbredning. Entomologisk Tidskrift (Stockholm), 58:
31-38.

1945. Foérteckning 6ver de av Zetterstedt i Insecta Lapponica
och Diptera Scandinaviae beskrivna tachiniderna med
synonymer jamte anteckningar Over en del arter.
Opuscula Entomologica (Lund, Sweden), 10: 26-35.

Robert, Adrien

1962. Les parasites et commensaux des nids de l’hirondelle
a front blanc, Petrochelidon pyrrhonota, au Lac Monroe,
parc du Mont Tremblant. Annals of the Entomological
Society of Quebec, 7: 87-92 [P. sialia, Quebec, cliff
swallow; not verified].

Rodhain, J., and J. Bequaert
1916. Histoire de "Passeromyia heterochaeta" Villen. et de

"Stasisia (Cordylobia) rodhaini" Ged. Bulletin
Scientifique de la France et de la Belgique, séries 7, 49:
236-289.

Rognes, Knut
1985. Revision of the bird-parasitic blowfly genus
Trypocalliphora Peus, 1960 (Diptera: Calliphoridae).
Entomologica Scandinavica, 15: 371-382, 23 text figs.
Rohdendorf, E. B.
1957. [On the parasitic flies, injurious to nestling singing
birds](In Russian, English summary). Entomologicheskoe
Obozrenie, 36: 116-124.

258 Protocalliphora

Rossi, F. W.

1848. Systematisches Verzeichniss der zweifligelichten
Insecten (Diptera) des Erzherzogthumes Osterreich. iii-x
+ 86 pp. Wien. [rearing of Musca azurea].

Sabrosky, Curtis W.

1950. Date of publication of Hall’s "Blowflies of North
America". Proceedings of the Entomological Society of
Washington, 52: 204.

1955. The taxonomic status of the armyworm parasite known
as Archytas piliventris (Van der Wulp)(Diptera:
Larvaevoridae). Florida Entomologist, 38: 77-83.

1956. The nomenclature of Protocalliphora (Diptera:
Calliphoridae). Proceedings of the Royal Entomological
Society of London, Series B, 25: 175-179.

1967. Corrections to A Catalog of the Diptera of America
North of Mexico. Bulletin of the Entomological Society
of America, 13: 115-125. [Trypocalliphora Peus 1960,
which included Protocalliphora hirudo (now braueri),
accepted as subgenus of Protocalliphora}.

1983. A synopsis of the world species of Desmometopa Loew
(Diptera, Milichiidae). Contributions of the American
Entomological Institute, 19(8): 1-69.

1984. A proposed amendment to Article 70b of the
International Code of Zoological Nomenclature on
misidentified type species. Bulletin of Foloeies
Nomenclature, 41(3): 152-155.

1989. Protocalliphora Hough, 1899 (Insecta, Diptera) and its
type species Musca azurea Fallén, 1817: Proposed
conservation of usage by designation of a replacement
lectotype. Bulletin of Zoological Nomenclature
46(2): 126-128.

Sabrosky, Curtis W., and Gordon F. Bennett

1958. The utilization of morphological, ecological, and life
history evidence in the classification of Protocalliphora
(Diptera, Calliphoridae). Proceedings of the Tenth
International Congress of Entomology (Montreal, 1956),
1: 163-164.

1971. The bird blow flies (Protocalliphora): Taxonomy,
biology and distribution. (Abstract). Proc. XIII
International Congress of Entomology (Moscow, 1968), 1:
194.

References 259

Sailer, Reece I., and Siegfried E. Lienk
1951. Blow flies (Calliphoridae) in Alaska. Canadian
Entomologist, 83: 208-211. [Apaulina sapphira, Tok,
Alaska, "cliff swallow" (error for bank swallow: Sailer in
litt.); = P. chrysorrhoea].
Sargent, William Dunlap
1938. Nest parasitism of hawks. Auk, 55: 82-84. [P.
splendida sialis (sic), Ithaca, N.Y., maggots in ears;
undoubtedly P. avium].
Schumann, H.
1986. Family Calliphoridae, pp. 11-58, in Sod(ed.), Catalogue
of Palaearctic Diptera, Vol. 12: 265 pp. Budapest.
Séguy, Eugene
1928. Etudes sur les mouches parasites. Tome I. Conopides,
OEstrides et Calliphorines de l’Europe occidentale.
Encyclopédie Entomologique, Sér. A, 9: 251 pp., 6 plates.
1929. Etude sur les diptéres a larves commensales ou
parasites des oiseaux de _ Jl’Europe _ occidentale.
Encyclopédie Entomologique, Sér. B, II. Diptera, 5:
63-82.
1955. Introduction a l’étude biologique et morphologique des
insectes diptéres. Publicagdes Avulsas Museu Nacional
Rio de Janeiro, 17: 1-260. [Protocalliphora; general

notes].
Seidensticker, John C., IV, and Harry V. Reynolds III
1971. The nesting, reproductive performance, and

chlorinated hydrocarbon residues in the red-tailed hawk
and great horned owl in south-central Montana. Wilson
Bulletin, 83: 408-418. [Protocalliphora sp., on hawks;
probably asiovora].

Shannon, Raymond C.

1923. Genera of Nearctic Calliphoridae, blowflies, with
revision of the Calliphorini. Insecutor Inscitiae
Menstruus, 11: 101-118.

1926. Synopsis of the American Calliphoridae (Diptera).
Proceedings of the Entomological Society of Washington,
28: 115-139.

Shannon, Raymond C., and Irene D. Dobroscky

1924. The North American bird parasites of the genus
Protocalliphora (Calliphoridae, Diptera). Journal of the
Washington Academy of Sciences, 14: 247-253.
[Numerous records, referred to under the various species].

260 _‘ Protocalliphora

Shewell, G. E.

1987. Calliphoridae. pp. 1133-1145, in J. F. McAlpine et al.,

Manual of Nearctic Diptera, Vol. 2.
Somme, L.

1961. On the overwintering of house flies (Musca domestica
L.) and stable flies (Stomoxys calcitrans (L.)) in Norway.
Norsk Entomologisk Tidsskrift, 11: 192-223.
[Protocalliphora azurea overwintering in crevices in
mountain huts].

Stabler, Robert M.

1959. Nesting of the blue grosbeak in Colorado. Condor, 61:

46-48. [P. metallica; det. Sabrosky, M. T. James].
Stein, Paul

1924. Die verbreitetsten Tachiniden Mitteleuropas nach ihren
Gattungen und Arten. Archiv fiir Naturgeschichte,
Abteilung A, 90(6): 1-271.

Stiner, Frederic M., Jr.

1969. Overwintering by Protocalliphora metallica (Diptera:
Calliphoridae). Annals of the Entomological Society of
America, 62: 1205-1206. [Newark, Del.; det. Sabrosky].

Stoner, Dayton

1926. Observations and banding notes on the bank
swallow--II. Auk, 43: 198-213. [Protocalliphorasp., Lake
Okoboji, Iowa; one example checked = hirundo).

1936. Studies on the bank swallow, Riparia riparia riparia
(Linnaeus) in the Oneida Lake region. Annals of the
Roosevelt Wild Life Forest Experiment Station of the
New York State College of Forestry at Syracuse
University, 4: 126-233. [P. splendida; not checked].

Storer, Tracy I.

1929. Protocalliphora in the nest of a mountain chickadee.
Condor, 31: 227. [P. splendida var. near hirundo, San
Bernardino Mts., Calif.; material apparently lost, possibly
= P, parorum Nn. sp.]

Taverner, P. A.

1934. Birds of Canada. Bulletin of the Canadian National

Museum, 72: 445 pp.
Theowald, Br.

1955. Calliphorinae (Dipt. Brach.) uit Vogelnesten.

Natuurhistorisch Maandblad, 44: 59-62.

References 26]

Thomas, Edward S.

1936. Additional records of Protocalliphora. Bird-Banding,
7: 46-47. [Protocalliphora spp., Ohio, 5 hosts; = P.
deceptor from Carolina wren and prairie and worm-eating
warblers, metallica from yellow-breasted chat and
Mississippi song sparrow].

Thompson, Patrick H.

1966. Arthropods from nests of the house sparrow.
Proceedings of the Entomological Society of Washington,
68: 44-48. [Protocalliphora larvae, Madison, Wis.].

Tirrell, Peter B.

1978. Protocalliphora avium (Diptera) myiasis in great
horned owls, red-tailed hawks, and Swainson’s hawks in
North Dakota. Raptor Research, 12 (1/2): 21-27, 2 figs.
[det. Sabrosky].

Townsend, Charles H. T.

1919. Description of the new species of Phormia. Annals of
the Entomological Society of America, 12: 379-380
(Appended to paper by Plath 1919c). [Phormia metallica
n. sp., holotype N.H.; paratypes are a mixture of
Protocalliphora metallica, P. aenea, and P. braueri].

1931. Notes on Old-World OEstromuscoid types, Part I.
Annals and Magazine of Natural History, series 10, 8:
369-391. [Notes on "holotype" of Musca azurea Fallén in
Stockholm].

1935. Manual of Myiology. Part II, Muscoid classification
and habits. Charles Townsend & Filhos, Itaquaquecetuba,
Sao Paulo, Brasil, 296 pp., 9 plates.

Van Tyne, J., and A. J. Berger

1976. Fundamentals of ornithology (ed. 2). John Wiley &

Sons, New York, xviii + 808 pp.
Verbeek, Nicolaas A. M.

1967. Breeding biology and ecology of the horned lark in
alpine tundra. Wilson Bulletin, 79: 208-218, 2 figs.
("blowfly larvae," Park Co., Wyo. = P. spatulata}.

1970. Breeding ecology of the water pipit. Auk, 87:
425-451, 6 figs. ["blowfly larvae," Park Co., Wyo.,
apparently P. spatulata].

Villeneuve, Joseph

1911. Notes diptérologiques. Wiener Entomologische
Zeitung, 30: 84-87.

1918. Sur Phormia sordida Zett. [Dipt.]. Bulletin de la
Société entomologique de France 1918 (10): 158-159.

262 — Protocalliphora

1928. Quelques mots sur les Calliphorinae paléarctiques.
Bulletin & Annales de la Société Entomologique de
Belgique, 68: 147-151. [Protocalliphora a subgenus of
Phormia; notes on confusion between European species
Phormia azurea Fallén and Phormia sordida Zetterstedt].

1931. Apercgus critiques sur le mémoire de P. Stein: "Die
verbreitetsten Tachiniden Mitteleuropas." Konowia, 10:
47-74. [Reiterates opinion that Protocalliphora can only
be a subgenus of Phormia].

Walsh, B. D.

1866a. Answers to correspondents: Rev. Jas. B. Fisher, N.Y.
Practical Entomologist, 1(10): 102. July 30. [Larvae found
attached to head and body of young swallow, identified
as "“Oestrus family" but undoubtedly larvae of
Protocalliphora, cf. Walsh 1866b].

Walsh, B. D.

1866b. Answers to correspondents: Rev. Jas. B. Fisher, N.Y.
Practical Entomologist, 2(1): 8. October. [Larvae from
Walsh 1866a identified by Osten Sacken as of "Musca
family in the vicinity of Musca or Sarcophaga."
Undoubtedly Protocalliphora}.

Walton, William R.

1914. Report on some parasitic and predaceous Diptera from
northeastern New Mexico. Proceedings of the United
States National Museum, 48: 171-186. [P. chrysorrhaea
(sic), horned lark; part = P. braueri, part in type series of
P. seminuda].

Wetherbee, Mrs. Kenneth B.

1932a. Two pairs of tree swallows mated during two
successive seasons. Bird-Banding, 3: 72-73. [P. splendida
var. sialia, Pomfret, Conn.; P. sialia, Worcester, Mass.,
pupae only].

1932b. Nesting bluebirds successful in spite of
Protocalliphora. Bird-Banding, 3: 73-74. [P. sialia,
Worcester, Mass., and Conn.; verified].

White, Clayton M.

1963. Botulism and myiasis as mortality factors in falcons.
Condor, 65: 442-3. [Calliphora sp., in heads of prairie
falcons, NW. Utah; possibly P. asiovora].

Whitehead, W. E.

1933. Parasites from a bird’s nest. Journal of Economic
Entomology, 26: 292-293. [P. splendida, Macdonald
College, Que., purple martin; not checked].

References 263

Whiting, A. R.

1967. The biology of the parasitic wasp Mormoniella
vitripennis [= Nasonia brevicornis] (Walker). Quarterly
Review of Biology, 42: 333-406, 45 figs.

Whitworth, Terry L.

1971. A study of the biology of the species of
Protocalliphora in the northern Wasatch Range. Utah
State University, Logan, Utah, unpub. MLS. thesis, 58 pp.

1976. Host and habitat preferences, life history,
pathogenicity and population regulation in species of
Protocalliphora Hough (Diptera: Calliphoridae). Utah
State University, Logan, Utah, unpub. PhD. dissertation
pp. xii + 144.

1977. [same title as Whitworth 1976]. Dissertation Abstracts

International, B, 37 (10): 4933-B.
Wing, Leonard W.

1956. Natural history of birds. A guide to ornithology.

Ronald Press Co., New York, xi + 539 pp.
Wood, E. D.

1953. Observations and report of bird-banding in the
vicinity of Victoria, B.C. Victoria Naturalist, 9: 76-78.
[Protocalliphora sp. or spp.; = probably P. braueri, at least
in part].

Woods, H. E.

1926. A new host for the blood-sucking larva fly,
Protocalliphora splendida. Bulletin of the Northeastern
Bird-Banding Association, 2: 12-13. [Huntington, Mass.,
barn swallow; = P. hirundo].

Zeleny, Lawrence

1968. Bluebirds for posterity. National Association for the
Protection and Propagation of the Purple Martins and
Bluebirds of America, Crawfordsville, Ind., 16 pp., 2 figs.
[Popular brochure; Apaulina sialia}.

1970. Most serious bluebird parasite is the blowfly. Purple
Martin Capital News, July 29, 1970: 12. [Popular article
in a commercial publication; P. sialia, det. Sabrosky].

1976. The bluebird. Indiana University Press, Bloomington,
Ind., pp. xix + 170. [P. sialia, etc., biology and effect on
birds].

Zetterstedt, Johann W.

1838. Diptera. pp. 477-868, in Zetterstedt, 1838-1840,

Insecta Lapponica, Lipsiae [= Leipzig], 1140 pp.

264 ~ Protocalliphora

Zumpt, F.

1956. Calliphorinae. Fam. 641i, 140 pp., 10 plates, in
Lindner, E. (editor), Die Fliegen der palaearktischen
Region. E. Schweizerbart’sche Verlagsbuchhandlung,
Stuttgart. Lieferungen 190, 191, 193.

1965. Myiasis in man and animals in the Old World.
Butterworth & Co., London, England, pp. xv + 267, 1
plate. [Protocalliphora, 9 spp. summarized].

Index to Bird Hosts 265

Index to Bird Hosts

blackbird, Brewer’s,103,127,128,132,
180,190,202,214,231,240
red-winged, 159,160,179,180,189,
190,206,207 ,231,251
yellow-headed, 128,175,176,179,
180,231
bluebird, eastern, 24, 176, 207, 214,
231
mountain, 214,231
unidentified, 22,28,29,42,127,128,
132,155,175,176,212,240,247,
248 ,250-252,254,262,263
western, 214,231
budgerigar, 31,32,34,36,127,128,231
bunting, indigo, 190,232
lazuli, 190,232
snow, 46,226-228,232
bushtit, 170,171,232
cardinal, northern, 155,156,190,191,
232
catbird, gray, 109-111,190,206,207,232
chat, yellow-breasted, 190,232,261
chick or chicken (domestic), 31,32,34,
36,122,128,231,247
chickadee, black-capped, 195,196,207,
232
Carolina, 155-157,232
chestnut-backed, 24,193,195,196,
232,245
mountain, 127,128,193,195-197,
214,232,245 ,260
unidentified, 22,28,36,80,84,
195,197
cowbird, brown-headed, 156,207,232
creeper, brown, 195,196,232
crow, American, 5,24,27,28,30,36,73,
103,105,108-110,155,156,214,
232,249
common = American
cuckoo, black-billed, 190,232
dipper, American, 96,97,232
dove, mourning, 40,102,103,233
ringed turtle-, 32,34,214,231
unidentified, 151
eagle, bald, 29,106,109,110,112,233,
241
golden, 37,100,102,103,105,122,
127,128,233 ,248,256
falcon, gyrfalcon, 106,109,110,233,
257
peregrine, 29,106,109,110,233,246,
247
prairie, 100,233,262

finch, Cassin’s, 167,233
house, 37,102,103,128,130,137,
150,151,167,170,171,233
purple, 233
rosy, 219,220,233
flicker, common = northern
northern, 233
red-shafted, 34,183,184,233
unidentified, 231
yellow-shafted, 214,233
flycatcher, alder, 119,120,127,129,131,
190,233
ash-throated, 214,233
dusky, 151,170,171,233
greatcrested,127-129,131,156,214,
233
Traill’s = alder, willow
unidentified, 36,150
western, 150,151,171,233
willow, 150,151,233
goldfinch, American,120,128,150,151,
190,207,234
lesser, 234
unidentified, 126,127,150,170,171
goshawk, see hawk
grackle, common, 36,96,97,120,127,
129,155,156,160,176,190,206,
207 ,213-215,234
grosbeak, black-headed, 127,128,133,
136,234
blue, 190,234,260
Pine, 162,163,234
gyrfalcon, see falcon
harrier, northern, 110,111,234
hawk, broad-winged, 29,109,110,234,
242
Cooper’s22,26,29,103,105,110,111,
234,242
ferruginous, 103,234
marsh = northern harrier
northern goshawk,103,109,110,
184,234
red-shouldered 29,110,111,234,240
red-tailed, 23,29,103,109,110,112,
183,184 ,234,252,259,261
sparrow = American kestrel
Swainson’s, 103,109,110,112,234,
261
unidentified, 5,25,28,29,73,105,
106,109-111,246,254,259
jay, blue, 207,234
Canada = gray
gray, 37,167,207,234

266 — Protocalliphora

pinyon, 103,234
junco, dark-eyed, 234
Oregon,34,128,196,234
slate-colored, 190,223,234
unidentified, 127,128,206,207,231
kestrel, American, 26,100.103,105,
183,184,214,233,248
kingbird, Arkansas = western
Cassin’s, 235
eastern, 119,120,150,151,190,
207,214,235
unidentified, 36
western, 127,128,150,151,214,235
lark, horned, 50,126-129,202,217,220,
235,261,262
magpie, black- billed, 21,28,30,102,103,
128,235,249,251,252,257
martin, purple, 29,176,214 ,235,239,
253,254,262
sand = bank swallow
meadowlark, eastern, 235
unidentified, 129,133
western, 127,128,235
mockingbird, northern, 156,235
nuthatch, brown-headed, 15,235,255
pygmy, 22,235,252
red-breasted, 195,196,235
unidentified, 22
white-breasted, 214,235
osprey, 110,235
ouzel,water=American dipper
ovenbird, 127-129, 235
owl, eastern screech-, 214,235
flammulated, 214,235
great horned, 102,103,105,106,109,
110,112,214,235,241,252,259,261
long-eared, 101,103,235
unidentified, 106
western screech-, 214,235
pewee, western wood-, 150,151,236
phoebe, black, 162,163,236
eastern, 96,97,155,156,176,214,236
Say’s, 96,97,162,163,214,236
unidentified, 36,96,97,162,241
pipit, water, 217,220,236,261
raven, common, 30,103,214,236
redstart, American, 117,119,120,121,
223,236,250
western=American
robin, American, 36,40,96,97,102,103,
108-110,126-129,150-152,167,
188,190,203 ,206,207,214,236,
250,251,256
screech-owl, see owl
shrike, loggerhead, 102,103,127-129,
131,236

siskin, pine, 128,132,236
solitaire, Townsend’s, 128,236
sparrow, American tree, 127,128,236
chipping, 127-131,139,141,167,
190,236
English = house
field, 190,236
fox, 190,236
grasshopper, 190,236
house, 127,128,130,132,136,145,
214,236,246,253,261
savannah, 30,127,129,130,190,226,
227,236,240
song, 127,128,179,180,190,207,
214,236,251,254,261
swamp, 159,160,190,236
unidentified, 30,36,73,126-131,
136,137,190,250
vesper, 127-129,131,190,236
white-crowned, 127,129,237
white-throated, 40,160,190,
206,207,237
starling, European, 43,102,103,128,176,
214,237,253
swallow, bank, 19,23,28,36,43,46,48,78,
127,129,142,144-147,172,175,
176,190,200, 214,237 ,240,252,255,
259,260
barn, 27,29,30,33,36,40,96,
97,102,103,116,127-129,131,133,
144,145,150,151,162,163,167,175,
176,195 ,196,202,214,217,237,
240,250,252,263
cliff, 27,36,103,127,128,137,142,
144,145,174-176,200,214,237,
242,257,259
eave = cliff
northern rough-winged, 160,190,
214,237
tree, 24,28,120,127,128,132,176,
190,207,214,237,240,242,250,
251,262
unidentified, 1,33,103,110,115,116,
163,167,175 ,215,241,247,262
violet-green, 127,128,131,144,
145,176,214,237
tanager, scarlet, 190,237
thrasher, brown, 190,207,212,237,251
thrush, hermit, 206,207,237
olive-backed = Swainson’s
russet-backed = Swainson’s
Swainson’s, 127,128,237
unidentified, 167
wood, 120,155,156,207,237
tit, great, 22

towhee, brown, 184,237
rufous-sided, 190,237
turtle-dove, ringed, see dove
vireo, blue-headed = solitary
red-eyed, 120,237
solitary, 117,119,120-122, 237,250
warbling, 223,237
warbler, Audubon’s, 34,151,238
black-throated blue, 117,119,120,
237
Canada, 127-129,237
chestnut-sided, 117,119,120,122,
238,250
myrtle, 119,120,122,127-129,207,
238
Nashville, 190,238
pileolated = Wilson’s
prairie, 155,156,190,238,255,261
unidentified, 36,73,126-128
Wilsen’s, 22,30,127,128,130,133,
238,257
worm-eating, 155,156,238,261
yellow, 30,119,120,127-129,
137,150-152,170,171,190,223,
238,254
yellow-rumped, 151,231,238

Index to Bird Hosts 267

waterthrush, Louisiana, 96,238
northern, 190,238
waxwing, cedar, 207,238
wood-pewee, see pewee
woodpecker, downy, 160,214,238
wren, Bewick’s, 196,238
Carolina, 155,156,238,252,261
house, 96,97,127-129,131,155,
156,196 ,207,214,238
long-billed marsh = marsh
marsh, 128,180,238
unidentified, 28,127,131,190
yellowthroat, common, 159,160,
190,238

268 — Protocalliphora

Index to Protocalliphora and Miscellaneous Names

(Family-group names are in capitals, valid genus-group names in boldface when
cited alone, valid species-group names in roman, and all synonyms in italics.

Specific names standing alone are in Protocalliphora, or have been at some

time.)

aenea, 4,6,14,30,50,51,58,59,63,
67,69,74,76,81,83,85,89,91,
93-99,129,135-137,148,150,160,
164,181,188,208,261

alcaeus, Pediobius, 37

amblyogma, 17

A paulina, 14,44-49,52,63,
171,246,248,259

Archytas, 62,258

armata, Morodora, 37,244

asiatica, 17,48

asiovora, 4,23,33,37,43,46,50,
58,59,66,67,74,77,82,88,
91,99-105,106,110,113,116,
154,157,244,248,249,251,
252 ,254,257,259,262

Avihos pita, 13,44,51,239

avium, 4-6,13,14,21-23,26,33,
36,37,43,44,46,50,58-60,63,
66,67,69,77,82,87,90,92,99,
100,101,103,105-113,116,147,
153-155,157,177,185,214,215,
217,221,240-244, 246,247,249,
251,254,257,259,261

azurea, 1,9-13,15-19,21,22,44,
46,48,68,105,122,129,137,
142,146,147,164,168,185,
224,239,242-245,248,253,254,
256,258,260-262

basingeri, 100,101,254

beameri, 4,57-59,68,79,80,84,88,
113-117, 248

bicolor, 4,21,58,59,65,66,68,76,
82,86,89,93,117-122,152,157,
185,192,197,224,244,249,250

Boreellus, 52,54,55

braueri, 4,5,13,14,17,19,22,23,25,
28,30-32,36,37,43,45-53,56,
58-60,62-65,76,81,84,88,90,
93,94,96,99,104,110,122-139,
146,154,157,168,192,193,202,
203,215 ,223,239,240,242-244,
246 ,247,249,250,254,256-258,
261-263

brunneisquama, 4,50,58,59,65,68,
78,81,85,116,139-141,167,248

caerulea, 12,18

Calliphora, 13,54,100,141

CALLIPHORINAE, 55

carnarida, 18

Carnus, 1,252

cavus, Dibrachys, 37

CHALCIDOIDEA, 37

Chrysomya, 16

CHRYSOMYINAE,-INI, 55

chrysorrhoea,-aea,-ea, 4,9,13,14,
16,17,19,23,36,37,44,46-48,50,51,
55,58,59,62,63,65,67,78,83,86,
90,92,113,116,122,126,129,141-
148,174-177,199,200,202,208,212,
217,221,240,255,259,262

Cochliomyia, 52

compacta, 17,51

Compsomyia, 105,242

CoOrusca, 18

cuprea, 122,123,126,130,137,188,
247,249,250,256

cuprina, 4,50,58,59,66,67,77,82,86,90,
93,96,121,148-152,163,164,167,
168,171,172,185,192,197,223,
224,251,254,256

cuprina var., 76,86,152

Cynomya, 185

Cynomyopsis, 54

deceptor, 4,50,53,58,59,63,65,67,77,
82,86,89,91,104,113,135,153-158,
192,208,241,245

Dibrachys, 37

dispar, 1,17,19,68

distincta, 17

ENCYRTIDAE, 37

EULOPHIDAE, 37

falcozi, 17

fallisi, 4,58,59,65,68,80,84,87,89,
91,99, 158-160,164,181,196,
203,227

frontosa, 17

gigantia, -tica, 181

groenlandica, Protophormia, 10

Haemoproteus, 26,27

halli, 4,33,50,58,59,67,78,82,88,
89,91,137,160-164,168,171,172,
202 224,227,228

Halteridium, 27

hesperia, 4,5,37,46,48,50,58,59,62,
67,78,82,85,88,122,130,136,
139-141,150,163,164-168,170-
172,203,246,249,254,256

Index to Protocalliphora and Misc. Names 269

hesperioides, 4,58,59,67,78,81,85,
89,92,141,163,164,166,168-172,
203,215,256

hirudo, 5,13,14,19,22,30,32,48,
49,51-53,60,93,122,123,125,
126,129-131,133,135,137-139,
188,239,240,242,244,247,254,
257,258

hirundo, 4,14,27,58-60,67,79,83,
88,90,92,100,113,116,147,172-
177,185 ,193,197,203,212,215,217,
252,260,263

interrupta, 4,58,59,65,68,79,84,88,
91,104,116,177-181,192,193

isochroa, 17

kaszabi, 17

lata, 4,47,48,50,58,59,65,67,69,
77 ,81,85,89,92,141,181-185

Leucocytozoon, 26

Lie i7,

lindneri, 17,19,44,48,49,51,138,
246,249

macellaria,Compsomyia, 105,242

maruyamensis, 17,45

megacephala,Chrysomya, 16

metallica, 4,13,14,21,33,36,46,47,
49,52,56,58-60,65 ,66,68,76,77,
79,84,86,89,91,93,103,111,117,
119,121,122,126,137,148,150-
152,157,160,180,181,185-193,199,
203 ,208,209,212,219,224,228,
229,243 ,244,249-251,253-255,256,
260,261

Mormoniella , 37

Morodora , 37,244

mortuorum, Cynomya, 185

Muscidifurax, 37

Mystacinobia

Nasonia, 37,41,157

Neottiophilum, 2

nidicola, 18

nigripal pis, - pus, 18

nuortevai, 17,228,246

Orneocalliphora, 44,48,142

parorum, 4,36,58,59,68,80,84,87,89,
93,121,180,181,193-197,221,
245,260

parva, 60,122,123,126,129

Pediobius, 37

peusi, 17

Philornis, 2,46

Phormia, 10,12,13,16,52,54,55,
245,261,262

PHORMIINI, 53-55,245

Plasmodium, 26

POLLENIINAE, 55

Protocalliphora, 44,53,55 299,300

PROTOCALLIPHORINI, 54

Protophormia, 9-13,18,21,52,54,55

proxima, 17

PTEROMALIDAE, 37

raptor, Muscidifurax, 37

regina, Phormia, 16,21,27,52,123,
245,251,254

reviso, 18

sabroskyi, 17

sapphira, 4,46-48,58,59,62,63,68,78-
80,84,86,87,142,144,146,197-200,

212,259

seminuda, 4,50,58,59,68,80,84,88,
90,92,126,129,172,200-203,224,
248,262

sericata, Phaenicia, 27,254

shannoni, 4,14,21,58,59,67,79,83,
87,90,92,157,177,185,192,203-
208,209,212,215,241,249-251,253

sialia, 4,6,14,22,23,33,36,37,47,58-
60,67,74,79,83,87,88,91,106,113,
116,147,171,176,177,185,192,197,
198,203 ,204,207,208-217,219,221,
240,241,243,247,249-255,257,259,
262,263

SIMULIIDAE, 65,243

sordida, 16-18,261,262

spatulata, 4,22,58,59,68,80,84,86,90,
92,192,199,212,217-221,244,261

spenceri, 4,58,59,68,79,84,85,87,90,92,
116,152,179,181,203,221-224

splendida, Calliphora, 13,30,93,96,
106,117,120-122,130,136,137,164,
171,172,175,185,186,188,191-193,
197,203,209,212,217,228-229, 239,
240,243,247 ,249-254,256,259,260,
262,263

surcoufi, 17

Tachinaephagus, 37,245

terraenovae, Protophormia, 9-13,
18,21

Trypocalliphora, 17,19,44,49,50,51,
53,63, 76,81,84,88,90,123,223,
246,249,257,258

tundrae, 4,16,46,48,58,59,62,65,67,
68,78,79,83,86,87,89,91,99,160,
164,181,196,224-228, 253-255

violacea, 17,18

viridescens, Calliphora, 141

vitripennis, Nasonia
or Mormoniella, 37,41,157,263

zealandicus, Tachinaephagus, 37,245

270 _—~ Protocalliphora

Figures

ocellar tubercle

median ocellus
preocellar area

frontal vitta

parafrontal

lunule

antenna
facial ridge

epistoma

prevertical
preocellar
area

Fig. 1. Head of Protocalliphora female, frontal view. Bristles:
fr, frontal; i vt, inner vertical; oc, ocellar; o vt, outer vertical; pc
0, proclinate orbital; poc, postocellar; pocl, postocular; rc o,
reclinate orbital; vb, vibrissa.

Fig. 2. Head of Protocalliphora male(a, frontal view, b, side
view), showing the measures used. ml, width of frons at
narrowest part; m2, maximum width of head (measured from
dorsal view); m3, maximum length of eye; m4, ocellar span; m5,
width of parafacial (opposite parafrontal-parafacial angle); m6,
vibrissal interval; m7, height of cheek.

Figures 27]

scutum (presutural area)

transverse suture
notopleuron

scutum (postsutural area)

‘+______greater ampulla

postalar callus

upper calypter

5 scutellum

AVA
\)
| ci

Woes
; jap sctLe

I

stem vein

lower calypter

3b

postalar wall

E tympanic membrane
7_i # suprasquamal ridge

lower calypter

scutellum

Wing (ventral)

subcostal
sclerite

Fig. 3. Thorax of Protocalliphora (a, dorsal view, b, detail of
postalar and suprasquamal areas). Bristles: acr, acrostical; ac npl,
accessory notopleural; a npl, anterior notopleural; ap sctl, apical
scutellar; dc, dorsocentral; p npl, posterior notopleural; post ia,

postsutural intraalar.

Fig. 4. Base of wing of Protocalliphora. a, dorsal; b, ventral.

272 ~~ Protocalliphora

deceptor

avium
6a (Shite)
avium chrysorrhoea braueri

Fig. 5. Cerci of Protocalliphora, posterior view.

Fig. 6. Aedeagus, of Protocalliphora, side view.

Figures 273

ae

10
avium asiovora deceptor lata
14 . { 13 14 | ;
chrysorrhoea aenea shannoni hirundo sialia
16 ze 18 | 19 :
Cuprina hesperia hesperioides halli Draueri

Figs. 7-20. Left surstylus, in profile, of Protocalliphora spp.
(s, surstylus, and c, cercus, on figs. 7 and 20).

274 Protocalliphora

21 | 24 | } 26
metallica parorum Cues fallisi seminuda sapphira ~
PXTh 28 i

31 32
spatulata brunneisquama Reais a eens beameri spenceri

Figs. 21-32. Left surstylus, in profile, of Protocalliphora spp.

Figures 275

36
tundrae hirundo

39
metallica

40

bicolor braueri

Figs. 33-41. Dorsal view of heads of Protocalliphora males.

276 ~ Protocalliphora

oe

pS

Fig. 42. Protocalliphora aenea. a, b, posterior region (note
lack of mesostigmatal folds and plaques); c, ventral cuticle; d,
dorsal cuticle (note lack of prominent cuticular ridges in c and d).

Figures 277

Fig. 43. Protocalliphora asiovora. a, posterior region (note
stigmatal area nearly bisected with folds and plaques); b, dorsal
cuticle; c, d, ventral cuticle (note interruption of medial band on
midline).

278 Protocalliphora

Fig. 44. Protocalliphora avium. a, b, posterior region (note
marked separation of stigmatal plates by folds in b); c, ventral
cuticle; d, dorsal cuticle (note prominent cuticular ridges in c and
d).

Figures 279

Fig. 45. Protocalliphora bicolor. a, b, posterior region (note
almost complete bisection of the stigmatal area); c, ventral
cuticle; d, dorsal cuticle (note reduction of cuticular ridges in c
and d).

280 Protocalliphora

|

Fig. 46. Protocalliphora braueri. Western material: a, b,
posterior region; c, dorsal cuticle; d, ventral cuticle. Eastern
material: e, posterior region; f, ventral cuticle.

281

Figures

ee

Ls

*

Ce

*

wes
os

ao

ae

aus

ca
23
ae)
—
aes
iS
eo
=
oo
6S
a €
ray
atek
[4°]
o
_-
ve
sh
=
ee)
S$ .2
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3B
xs P
i)
SE
ae)
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~=

)

d

1c

Protocall
le;

c, dorsal cut

47.

Fig.
ridges in c and qd).

282 Protocalliphora

Fig. 48. Protocalliphora cuprina. a, b, posterior region; c,
dorsal cuticle; d, ventral cuticle.

Figures 283

Fig. 49. Protocalliphora deceptor. a-c, posterior region (note
very small spines and resulting fine-sandpaper appearance; also
note lack of circumstigmatal folds; d, dorsal cuticle; e, f, ventral
cuticle.

284 Protocalliphora

Fig. 50. Protocalliphora fallisi. a, posterior region; b, dorsal
cuticle; c, ventral cuticle (note much reduced ventral bands).

Fig. 51. Protocalliphora halli. a, b, posterior region; c, dorsal
cuticle; d, ventral cuticle (note marked reduction, almost absence,
of posterior band).

286 —- Protocalliphora

Fig. 52. Protocalliphora hesperioides. a, b, posterior region
(note bisection of stigmatal area by mesostigmatal folds and
plaques); c, dorsal cuticle; d, ventral cuticle (note reduction of
cuticular folds in c and d).

Figures 287

Fig. 53. Protocalliphora hirundo. a, b, posterior region; c,
dorsal cuticle; d, ventral cuticle (note heavy cuticular ridging in
c and d, with spines directed both anteriorly and posteriorly.

288 Protocalliphora

Fig. 54. Protocalliphora interrupta. a, posterior region; b,
dorsal cuticle; c, d, ventral cuticle (note interrupted medial
band).

Figures 289

Fig. 55. Protocalliphora lata. a, b, posterior region; c, dorsal
cuticle; d, ventral cuticle (note prominent cuticular folds on both
dorsal and ventral cuticle, and spines directed both anteriorly and
posteriorly).

290. ~~ Protocalliphora

Fig. 56. Protocalliphora metallica. a, b, posterior region; c,
dorsal cuticle; d, ventral cuticle.

Figures 291

Fig. 57. Protocalliphora parorum.
dorsal cuticle; d, ventral cuticle.

a, b, posterior

region; c,

292 ~~ Protocalliphora

Fig. 58. Protocalliphora seminuda. a, posterior region; b,
dorsal cuticle; c, ventral cuticle.

Figures 293

Fig. 59. Protocalliphora shannoni. Eastern material: a, b,
posterior region; c, dorsal cuticle; d, ventral cuticle.

294 Protocalliphora

Fig. 60. Protocalliphora shannoni. Western material:
posterior region; c, dorsal cuticle; d, ventral cuticle.

295

Figures

oO oO
oo
(=|
oe

he
o
oe
eo
ae)

poster

a,
icle (note pronounced cut

}

b

AS)
SS
=
ny —
nN
e078
aye!
foe
Ro 8
= aC
—~ & Ly
S5=-c
oe eg
OO 2
en =
Smee)
ee
ee
= ¢
Ses
mee
F re
OO
mW naw;v
u
og
O10)

296 Protocalliphora

Fig. 62. Protocalliphora spatulata.
dorsal cuticle; c, ventral cuticle.

a, posterior

Figures 297

Cc

Fig. 63. Protocalliphora spenceri. a, posterior region; b,
dorsal cuticle; c, ventral cuticle.

298 — Protocalliphora

Fig. 64. Protocalliphora tundrae. a, b, posterior region; c,
dorsal cuticle; d, ventral cuticle (note reduced posterior band).

Map |. Distribution of Protocalliphora in North America, as
reflected by collecting localities. Major centers of collecting are
starred, in addition to concentrations of localities in northern
Utah and California.

300 Protocalliphora

p lo
| es

| i

| | =~ = Ge 45
| | | >

| :

| :

s

| | ad

Map 2. Palearctic distribution of Protocalliphora, based on
known extreme southern localities and assumed northern limits
based on comparable latitudes in North America. Numerous
records are known for Europe, but few elsewhere in Eurasia.

Map 3. Distribution of Protocalliphora braueri in North
America.

302 Protocalliphora

ay he

va 2 N b if
7 Wh Y, S . S e
AA a VA | a : aN X
y Yi < ~~
ey a \

See EU Geer

Ds

N @
“To ©
Hen)
@ Sel
. gay
u

| pl

/

Map 4. Distribution of Protocalliphora aenea (stars), P.
beameri (small circles), and P. deceptor (large circles).

Maps 303

Map 5. Distribution of Protocalliphora avium (circles) and P.
asiovora (stars).

304 = Protocalliphora

Map 6. Distribution of Protocalliphora chrysorrhoea (circles)
and P. seminuda (stars). The records of chrysorrhoea for
Michigan and Minnesota are "probables," not verified.

Map 7. Distribution of Protocalliphora cuprina (circles) and
P. interrupta (stars).

306 = Protocalliphora

Map 8. Distribution of Protocalliphora halli (small circles),
P. hesperia (large circles), and P. hesperioides (stars).

Maps 307

Map 9. Distribution of Protocalliphora hirundo.

308 Protocalliphora

Map 10. Distribution of Protocalliphora lata (circles) and P.
parorum (stars).

Map 11. Distribution of Protocalliphora metallica (circles)
and P. bicolor (stars). The record of metallica for South Dakota
is by state only, without definite locality.

310 Protocalliphora

Map 12. Distribution of Protocalliphora shannoni (small
circles), P. spatulata (large circles), and P. tundrae (stars).

Map 13. Distribution of Protocalliphora sialia. The record
for South Dakota is by state only, without definite locality.

312 —Protocalliphora

Map 14. Distribution of Protocalliphora spenceri (circles) and
P. brunneisquama (stars).

ai) wy

ee ie
se

ees
ita

i
acces
i Shall Wy Pitie ee

t
Sia vale

The Calliphoridae are a widespread family of flies whose larvae are
obligatory bloodsucking parasites of nestling birds. Part of the Smithsonian
Research Monograph Series, this long-awaited study describes the North
American representatives of the blow fly genus Protocalliphora and
recognizes twenty-six North American species, including fifteen species
newly described.

‘This monograph is a work of genuine scholarship—the distillation of
knowledge of many years by acknowledged experts in the field. The
senior author is the dean of dipterologists, recognized internationally and
for decades for the high quality of his research. . . [This book] fills an
important gap in our knowledge of calliphorids of the genus Protocalli-
phora. It brings together scattered research and records on biology,
ecology, and distribution, and it resolves taxonomic problems and
inconsistencies, which will facilitate the study of these intriguing flies.”’

—Bernard Greenberg
Department of Biological Sciences
University of Illinois at Chicago

About the Authors

Curtis W. Sabrosky is a cooperating scientist at the Systematic Ento-

mology Laboratory, U.S. Department of Agriculture, and a research

associate at the National Museum of Natural History, Smithsonian

Institution. He is former president of the Society of Systematic Zoology,

the Entomological Society of America, and the Entomological Society of
Washington. Gordon F. Bennett is professor of biology at Memorial

University of Newfoundland and the head of the International Centre for

Avian Haematozoa at the university. Terry L. Whitworth is owner and .
_ president of Whitworth Pest Control, Inc., Tacoma, Washington.