Idea Transcript
BSOVE
11( 1):
ISSN
1- 198 ( 1986)
0146- 6429
Bulletin of the
SOCIETY OF VECTOR ECOLOGISTS,
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Volume 11, Number 1
September, 1986
BULLETIN OF THE
SOCIETY OF VECTOR ECOLOGISTS Volume 11 - Number 1 September 1986
James P. Webb, Ph.D., Editor
Orange County Vector Control District 13001 Garden Grove Boulevard Garden Grove, CA 92642
Minoo B. Madan, Production Manager California State Department of Health Services Los Angeles, CA 90026
EDITORIAL BOARD Dr. R. S. Lane
Dr. F. J. Radovaky California Acad. of Sciences
Liverpool, England
University of California Berkeley, California
Dr. A. Ali
Dr. M. S. Dhillon
Dr. L. A. Magnarelli
Dr. R. E. Ryckman
University
Northwest M.A.D.
Conn. Agr.
Sanford, Fiordia
Riverside, California
New Haven, Connecticut
Loma Linda, California
Dr. D. R. Barnard
Dr. S. P. Eisen Institute
Dr. C. J. Mitchell Zool. Institute Universitiit
Dr. H. L Scudder
Lone Star Tick Research Lab. Poteau, Okalhom
Antwerp, Bdgium
Zurich, Switzerland
Hayward, California
Dr. Norbert Becker
Dr. R. E. Fontaine
Dr. G. R. Mullen
Dr. M. W. Service
University of Heidelberg Heidelberg, West Germany
CDC, Public Health Service
Auburn
Liverpool Sch. of Trop. Med.
Atlanta, Georgia
Auburn, Alabama
Liverpool, England
Dr. H. Briegel University of Zurich
Dr. M. L Goff University of Hawaii, Manoa
Dr. S. Nalim
Dr. E. S. Tikasingh
Caribbean Epidemiology Ctr.
Zurich, Switzerland
Honolulu, Hawaii
Vector Control Research Unit Ungaran, Indonesia
Dr. P. Carnevale
Dr. D. J. Gubler CDC, Public Health Service San Juan, Puerto Rico
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Dr. G. B. White
Vero Beach, Florida
London, England
Dr. A. J. Adams' University of Panama
Dr. J. B. Davies Liverpool Sch. of
Panama
Florida
of
O.R.S.T.O. M. Bobo Diolasso, Burkina Faso
of
Trop. Med.
Trop.
Medicine
Kip.
Station
University
San Francisco, California
Loma Linda University
Cantonal* State University
Port of Spain, Trinidad
London Sch. Hyg. Trop. Med.
Dr. S. C. Williams
Dr. C. E. Hopla University of Oklahoma
Dr. S. Palchick
Fort Detrick, Maryland
Norman, Oklahoma
St. Paul, Minnesota
San Francisco State Univ. San Francisco, California
Dr. W. J. Crane Rutgers University
Dr. M. J. Klowden University of Idaho
M. A. Parsons
Dr. W. J. Wrenn
Moscow, Idaho
Columbus, Ohio
Dr. LA. Lacey Medical Services Consultants Arlington, Virginia
Texas Tech University
Dr. G. G. Clark U. S.
Army
Medical Res. Inst.
New Brunswick, New
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University
Upsala, Sweden
Jersey
Metropolitan
Department
Moeq.
Cont.
Health
of
Dr. D. B. Pence Lubbock, Texas
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BULLETIN OF THE
SOCIETY OF VECTOR ECOLOGISTS VOLUME 11
SEPTEMBER, 1986
NUMBER 1
CONTENTS In Memoriam - Richard B. Loomis( 1925- 1985)
iv
In Memoriam - James M. Brennan( 1905- 1985)
vii
Submitted Papers A Chigger
Evidence
Bibliography
1758- 1984 ( Acari: Trombiculidae): Systematics, Biology, and Ecology M. L. Goff, R. B. Loomis, and W. J. Wrenn
Incriminating Coquillettidia
perturbans (
Diptera: Culicidae) as an Epizootic Vector of
Eastern Equine Encephalitis. I. Isolation of FFF. Virus from Cq. perturbans During an Epizootic Horses in New Among
Evidence
Jersey
Incriminating
W. J. Crans
Coquillettidia
perturbans (
1
and
T. L. Schulze
178
Diptera: Culicidae) as an Epizootic Vector of
Eastern Equine Encephalitis. II. Ecological Investigations Following an Inland Epizootic in New
Jersey
W. J. Craps, L. J. McCuiston,
and
T. L. Schulze
185
Impact of Two Unicellular Algae on the Development of Culex quinquefasciatus Say ( Diptera:
Culicidae) Survival
L. S. Mian
and Ovipositional Response Sewage Effluent
of
Culex
quinquefasciatus
and
M. S. Mulla
191
Say ( Diptera: Culicidae) to L. S. Mian
M. S. Mulla
194
Field Tests in Western Montana of Mechanical Transmission of Tularemia by Biting Flies Diptera) Between Immobilized
197
and
Scientific Note
Laboratory
Animals.
C. B.
Philip
and
W. L. Jellison
4
NUMBER 1
SEPTEMBER, 1986
VOLUME 11
IN MEMORIAM
DR. RICHARD B. LOOMIS 1925 - 1985 mites as well as advanced knowledge about vertebrate
1985, Dr. Richard B. Loomis,
On October 8,
zoology, particularly the mammals and reptiles. Dick
Professor of Biology at California State University, Beach home at Long Beach( CSULB) died at his Long
60.
age
born in Lincoln,
Grant) at the University of Kansas Medical Center in
Professor Loomis had been
CSULB for 30
in
there as an Instructor
received a Research Fellowship ( Office of Naval Research Contract) early in his years ( 1948- 1953), and then a Research Assistantship ( NIH-USPHS
Biology
in
summers
was
and was an expert teenager
was a
and
Dick
1955.
Nebraska in 1925 the time he
beginning
years,
time
spare
associated with
herpetologist
having
on
field
spent
trips
Kansas
by
City(
1953- 1955).
From Lawrence, Dick and
his family moved to Long Beach where he continued
his early
his productive canter at CSULB as an instructor( 1955-
collecting
4'
F
tr
4, tt.....
.
4111111hik‘ 1
amphibians
and
reptiles.
Biology
was
a
latter days of
of World
1956),
strong
driving force in Dick' s life from a very early age until his dying day. After serving as a naval officer in the
(
interest
and
In 1955, he the
was awarded a
University
time
at
fundamental
of
education
acquired
about
1963- 1985).
California,
Ph.D.
Kansas ( KU).
KU that he
Assistant
Professor ( 1956- 1959),
and
Dick taught a multitude of courses
during his tenure at Long Beach State including General Biology, General Zoology, Animal Life in
War II, he attended the University where he earned a B. S. in
Nebraska ( Lincoln)
Zoology in 1948. in Zoology from was during this
an
Associate Professor ( 1959- 1963), and Full Professor
General
Entomology,
Field
Natural
History, Medical Entomology, Principles of Animal Taxonomy, and General and Advanced Herpetology. Many of Dick's graduate students were first introduced
It the
chigger
iv
NUMBER 1
to
the
SEPTEMBER, 1986 of
field
Herpetology
class.
wonders
Dick's
load Dick
also
biology In
while
addition to
enthusiastically
enrolled
in
( 1970- 1971),
his teaching
served
on
administrative and academic committees.
Foundation.
various
He
Committee, Committee,
pocket provided the backing for Dick's extensive field
Biology leadership Biology Department Advisory
collecting ventures. Through the years Dick actively
collected small vertebrates and their ectoparasites from
many parts of the world including California,
Curriculum
Committee, Research R.T.A.& P. Committee, Graduate Committee, Hiring Committee, New Committee, Campus Affairs Committee,
Studies
Building Scholarship& Loans Committee( Teaching Excellence Committee.
Chairman),
Colorado, Florida, Hawaii, Idaho, Illinois, Iowa, Louisiana, Maine, Minnesota, Mississippi, Missouri, Montana, Nebraska, Nevada, New Mexico, New York, North Carolina, North Dakota, Oklahoma, South Dakota, Tennessee, Texas, Utah, Virginia,
and the
Wisconsin, and Wyoming in the U. S., and Baja California ( Norte and Sur), the states of Cahuila, Chihuahua, Durango, Guerrero, Jalisco, Mexico,
Academic honors awarded to Dick also included, 1) Fellow of the Herpetologist League, 2) Fellow of the Southern California
Michoacan, Nayarit, Sinaloa, and Sonora in Mexico,
Academy of Sciences, 3) Phi Kappa Phi, 4) Sigma Xi, 5) CSULB Distinguished Scholar( 1982). He was also active in
and New Guinea, New Zealand, and Australia. Dick's field trips were always a highlight for
anyone fortunate enough to go along on one with him. These trips resulted in thousands of vertebrate
a number of professional
societies:
specimens and hundreds of thousands of ectoparasite
Southern California Academy of Sciences Vice- President and President, Chairman or CoChairman for the first five annual meetings); American Society Member( member
since
American
Society
of
since
under
1960);
Society
for the
Study
1960);
Society
member since
As
an
number of
avid
of
under
researcher,
and
since
was
awarded
helped
(
a
(
to support
himself in obtaining his own education and assisted many of his graduate students at CSULB. Most noteworthy is the NIH-USPHS Grant AI-03407, the
Chiggers of Western North America, the Principal Investigator from 1960 Other
grant
included
an
stipends
Office
that
were
that
he had to
Kethley.
Dick's
critical
and
fair
supervision.
Their
1966), Ronald E. Somerby ( 1966), Julius C. Geest 1966), Lynell K. Tanigoshi ( 1968), Richard M. Davis ( 1968), Jerry L. Fowler ( 1968), Norman G. Puckett ( 1969), James P. Webb, Jr., ( 1970), James L. Lucas ( 1970), W. Calvin Welboum ( 1972), M.
Lee Goff ( 1974), Lawrence V. Pomeroy ( 1974), Sherburn R. Sanborn ( 1977), Stephen G. Bennett
1977), Hans Megens( 1980), Charles S. Rau( 1980), Steven D. Werman ( 1980), Kazuhiro Ando ( 1983), Lawrence L. C. Jones( 1985), Noor S. Tietze( 1986), and
Gerald E. Greene ( 1986).
Of these individuals
whom we know personally, it can be said that we feel
as
that we belong to an elite fraternity that had the good fortune of having had Dick Loomis as our faithful counselor and friend during our stint at CSULB.
through 1974.
awarded
B.
program are included here: Marilyn Bunnell ( 1957),
of
Dick
Dr. John
Delmer E. Mangum ( 1958), Alvin H. DeYoung 1958), Kenneth Leith ( 1958), Charles M. Page 1961), William J. Wrenn ( 1965), W. Leon Hunter
Vector Ecologists
which
of
names and the year each finished his or her degree
1983).
Research Grants
curatorship
or the resulting data to write a masters degree thesis
of
Amphibians and Reptiles( Charter Member); Entomological Society of America( member since 1973); Acarological Society of America( Charter Member); Desert Protective Council( member since
the
Directly or indirectly, 27 individuals developed the background, format, and information from these trips
1947); American Southern
Society Parasitologists( and California Parasitologists) ( member of
collection ( ca.
vials) have recently been acquired by the Division of Insects, Field Museum of Natural History, Chicago,
1947); Kansas
Ichthyologists
Herpetologists ( member
chigger- mite
110,000 prepared microslides; ca. 12,000 alcohol
Academy of Science( member since 1948); Texas Academy of Science( member since 1970); Southwestern Association of Naturalists( member since 1969); Society Systematic Zoology( Charter Member);
Dick's
specimens.
Mammalogists, Life
of
and several Faculty Grants, CSULB
These grants and oftentimes money from his own
was
Graduate Coordinator for the Department of from 1965 to 1978. He provided input and as a member of the
VOLUME 11
him
Naval Research Contract, Kansas ( 1948- 1952), a NIH-USPHS
Dick's legacy to all of us was to develop our interests
School, Kansas Grant, CSULB
that all of Dick's former graduate students are
of
University of Grant, University of Kansas Medical City ( 1953- 1955), a U. S. Army
and potentials in the field of biology to such a degree gainfully employed in the scientific field in some v
least 12
or another and at
form
NUMBER 1
SEPTEMBER, 1986
VOLUME 11
of the
27
went on
WE MUST SAIL ON
to
obtain their doctoral degrees in biological disciplines. BSOVE is dedicated to Dick friend and Loomis and the late James M. Brennan, a first 177 The years. for Dick' s of colleague many 1758pages is comprised of," A Chigger Bibliography
This issue
1984 ( Acari: Trombiculidae): and
co- authored
Ecology, "
B. Loomis, more than
them
and
100
by M. Lee
during
with chigger mites.
bibliography
provide
Systematics,
To
and
the baseline
knowledge for future
his
We must sail on, he's disembarked
To some port so far away
We must sail on but we still can hope
of
included
We must sail on though were now becalmed And wait for the next strong wind
with others, will
they, along
nucleus
We'll see him again someday
career, most of
are
trombiculid
chartered shore
Though he will sail with us no more
Biology,
Goff, Richard
They
our own=
We must sail on in this wayward ship
William J. Wrenn. Dick co- authored
publications
had to do
in this
We must sail on in this ship of life
of the
And pray to God that a man like him
mite
Will sail with us again
researchers.
Dick is survived in body by his loving and patient wife
Margaret
of
Long
Beach, two daughters,
Written in memory of Richard B. Loomis
Orange, Linda Garrett of Long Beach, Julia Bynon of sister Beach, his his son Richard Loomis of Long
by Doug Bynon
Mary Margaret Moulton of Lincoln, Nebraska, and
seven grandchildren. In spirit Dick is survived by the scientific community at large, whom he loved and respected and who loved and respected him. James P. Webb, Jr.
Orange County Vector Control District Garden Grove, CA Stephen G. Bennett
Orange County Vector Control District Garden Grove, CA William J. Wrenn
University of North Dakota Grand Forks M. Lee Goff
University of Hawaii at Manoa Honolulu
vi
NUMBER 1
SEPTEMBER, 1986
VOLUME 11
IN MEMORIAM
DR. JAMES M. BRENNAN 1905 - 1985 r
g" f4
od
r ,..
r,,A0..-
,
1,,,
4' V0y,,,,,,,,,. ''''',.,,
406.,„,.„
4,,,,, k ti
o,,,,
ot,
v ,
.
4. ad
t
4r
i .
Dr. James M. Brennan died in Queen' s Medical on 31 October 1985 at the age of 80. He is
Center
survived
by his
wife
Dr. Brennan
BA
was
Isabel.
which
Pennsylvania. His from Dickinson College with a major in was a native of
English in 1926. He
received
his MA in 1931
and
Ph.D. in 1933 from the University of Kansas, majoring in entomology. Following his graduation, Dr. Brennan worked with the USDA Plant Quarantine from
1935
1941
to
entomologist
then
and
for the U. S.
as
a
civilian
from 1942 to 1944. In 1944, Dr. Brennan joined the U. S. Public Health
Service
Army
and spent most of
Mountain
Following
Laboratory his
Dr. Brennan
his
in
career at the
Hamilton,
Honorary Associate
in
Entomology
was a
Fellow
Association for the Advancement
Entomological
American Mosquito
Control
Systematic
Zoology.
While
Society
the
Montana.
from the USPHS in 1975, to Hawaii and was appointed an
Museum. Dr. Brennan member of the
Rocky
retirement
moved
the
at
of the
of
of
Association,
majority
of
and
1940' s
was
on repellents
involved in
for ticks
work
and a
South America.
While his
entomological activities were diverse in nature, Dr.
Brennan's major area of interest was in the taxonomy of the family Trombiculidae, or chiggers. During his research career, he was considered the major world
authority on the taxonomy of this difficult group of acarines.
assembled
At the Rocky Mountain Laboratory, he a reference
collection
which would
ultimately become the National Chigger Collection
of the U. S. National Museum of Natural History/ This collection, currently
world.
America,
Jim will always be fondly remembered by his colleagues and friends not only for his ability as an
of
acarologist and teacher, but also for this interest in
fishing and his delightful sense of humor. Acarology
more than
during
an.
Hawaii at Manoa, has become the largest single
is the poorer for this passing.
100 published works dealt with the systematics of the Trombiculidae, he
drugs in Central.
reference collection for this group of mites in the
American
Dr. Brennan' s
Also, during the 1940's, Dr.
housed at the Acarology Laboratory, University of
Bishop
Society
use.
Brennan was involved in the testing of antimalarial
Smithsonian Institution.
American
Science
Society
Parasitologists,
of
and in the 1952 USDA Yearbook, is credited for
development of the repellent issued to the military is still in
M. Lee Goff
the
and other arthropods;
University of Hawaii at Manoa, Honolulu vii
BULL. SOC. VECTOR ECOL., 11( 1):
1- 177.
SEPTEMBER, 1986
A CHIGGER BIBLIOGRAPHY 1758 - 1984
ACARI:
TROMBICULIDAE):
SYSTEMATICS, BIOLOGY, AND ECOLOGY! M. Lee Goff2, Richard B. Loomis3, and William J. Wrenn4
ABSTRACT: A comprehensive bibliography of publications dealing with the family Trombiculidae (including
the Leeuwenhoekiidae of some authors) from 1758 through 1984 is presented. These 2,441 references deal with
systematics, taxonomy, ecology, morphology, life cycles, histology, host-parasite relationships, physiology, behavior, and medical problems directly related to trombiculid mites. INTRODUCTION It
has
Wharton
and
missed.
We anticipate that additions and
corrections will be published
been nearly
comprehensive chigger
by
been
35
years
since
bibliography
was
Fuller ( 1952)
appropriate to publish another.
and
We have
last
the
completed
it
seems
attempted to
periodically.
To avoid
delay in publication, we have not indexed the references. We view this bibliography as a basis for future analyses and comprehensive listings of taxa and topics.
include all of the known references to the chiggers, family Trombiculidae ( including
Leeuwenhoeof Systema Naturae( 1758) through December, 1984. We included kiidae), from Linnaeus'
10th
the
edition
references to systematics, cycles,
histology,
ecology, morphology, life host- parasite relationships, physio-
logy, behavior, and medical problems which deal directly to
with
include
trombiculid mites.
We have
which
deal exclusively
siosis ( scrub
merely
typhus or
make
levels. Papers
chigger-borne
tsutsugamushi
reference
This
excluded.
with
to
includes
taxa or
chiggers
rickett-
disease) have
or
been
most texts and medical
many host-parasite lists, and such as Mammalian Species which are secondary without new information. Also excluded are
references, as well
reviews sources
title- only and most abstract references.
references
we
have
endeavored
up to 1985, it is
year.
Use of
Style Manual. Those publications not seen are indicated by an asterisk(*).
When the title is not in the language of the text, it
will
be explained;
e. g.,
a title in English of a
Russian publication with an English summary is stated
as (
in Russian with English summary).
Abbreviations of periodicals or words used conform as far as possible to the style in the Serial Sources for the Biosis Data Base, Volume 1984, or the International List of Periodical Title Word Abbreviations, ISO 833- 1974. However, titles of
certain journals ( such as some in Japanese, Chinese, and Russian) seem to have changed over time from the
Although
chronologically by
authors' initials follows the 5th edition of the CBE
attempted
all publications which propose new
recommend taxonomic changes on all
The citations are arranged alphabetically by
author( s) and then
to
include
certain that some
all
have
originals
translations.
or
adopted
different
English
We have used any title in English
supplied as part of an original publication.
Journal Series No. 3048 of the Hawaii Institute of Tropical Agriculture and Human Resources. This paper
is based on studies supported by NIH grant AL 20520 to the Department of Entomology, University of
Hawaii at Manoa and NSF grant DEB 7925107 to California State University, Long Beach.
2Department of Entomology, University of Hawaii at Manoa, 3050 Maile Way, Honolulu, Hawaii 96822, USA. 3Department of Biology, California State University, Long Beach, California 90840, USA (Deceased). 4Department
of
Biology, University
of
North Dakota, Grand Forks, North Dakota 58202, USA.
rr.
VOL 11, NO. 1
BULL. SOC. VECTOR ECOL.
2
As this
manuscript
was
being
authors( MLG, RBL, and WJW) felt it appropriate to dedicate this
M.
The authors extend special thanks to Noor S.
and colleague who contributed
Tietze for helping to assemble, correct, and standar-
bibliography
Brennan, a dear friend
to the late Dr. James
his impressive list of publications to trombiculid mites. Dr. Brennan passed
much more than
the
study
away
on
of
31 October 1985.
completion
of
the
Regrettably,
manuscript
came
the
prior
to
untimely
Dr. Richard B. Loomis, our teacher, mentor, passed away colleague, and dear friend. Dr.. Loomis
death
on
of
8 October 1985.
this
bibliography
such great
lives.
We ( MLG
and
WJW) dedicate
to both these individuals
impacts
Acknowledgments
the
assembled,
who
had
on our professional and personal
dine the references. We thank Cliff Hill and Jeffrey Alberts for their
bibliography
on
assistance
floppy
in
disks.
the
formatting We
also
extend
special thanks to Dawn Rath who prepared the final draft. We are grateful to the Department of Biology,
University of North Dakota and the Institute of Parasitology,
California
State
University,
Long
Beach, for providing assistance. In addition. we thank Dr. James P. Webb and Stephen G. Bennett for critical review of
draft
copies of this
bibliography.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
3
BIBLIOGRAPHY 0001 * Abussalam,
0002
Abonnenc,
1939.
E. &
larvaire
de
Abonnenc,
Ah,
d'
un
1: 83- 86.
Trombiculidae ( Acarien):
nouveau
Taufflieb. du
region
n.
E. &
Ann.
phacochere.
Sun.
1960. from
mite
1957b.
Ah,
Hyong-
Sun.
un nouveau
Entomol.
0008
Soc.
pilosa
Pathol.
n.
Exot.
gracilis,
( Acarina,
sp.
C. E. & J. Lipshutz. Pacific Theater. J. Am.
Alberti,
G.
1979.
and
Allison,
ed. ,
F. R.
50: 556- 67. a
new
species
Trombiculidae) .
0011
Utah.
0012
D. M.
Utah
Allred,
Utah.
trom-
1954.
Parasites
of
124: 1095- 1100.
Proc.
1958a.
Zealand Occas.
the
on
Utah Acad.
Sci.
New Zealand 2) : 419- 23.
reptiles.
Publ.
(
stylosome
(
feeding
tube)
of
31: 61- 63.
Mites found on mice of the genus Peromyscus in
infestation.
Families
New
Ser.
Observations
1956.
General
D. K.
Tsutsugamushi fever in the Southwest
Assoc.
organs of Actinotrichida. Pp. 501- 07, in J. G. Recent Advances in Acarology, Vol. II. Academic Pr. ,
chiggers.
IV.
1944. Med.
New Zealand Wildl.
D. M. I.
Allred,
of
Parasitology
Fine structure and probable function of genital
1982.
herpetology.
some
Trombicula
Claparede
New York.
Allred,
et
New intranasal chiggers of the genus Microbats in Korea ( Acarina: Trombiculidae). J. Med.
Ahlm,
Rodriguez,
0010
vercam-
1: 186- 91.
papillae
0009
some
Trombiculide
de Schoengastia radfordi Jadin et Vercam-
Shunsennia
Korea
1964.
from
trombicula
0007
Hum.
Deux trombiculides nouveaux de
Euschoengastia
n.
Hyong-
Sur
50: 577- 80. 0006
Euschoen-
Parasitol.
Euschoengastia ( Helenicula) Pathol. Exot. 50: 86- 89.
Soc.
Taufflieb.
Tchad,
sp. , presence Grandjean, 1954. Bull.
biculid
1957a.
Tchad:
Bull.
sp.
R.
du
region
youhensis
0005
un
parasite
R.
la
mengrandjeani
men-
Sur
Entomol.
30: 219- 24.
Abonnenc,
la
Indian J.
1955.
rouchoni,
Comp.
0004
A new trombiculid larva parasitic on the house
rattus).
E.
gastia
0003
M.
( Rattus
rat
Great Basin Nat.
16: 23- 31.
Mites found on mice of the genus Peromyscus in Laelaptidae
and
Phytoseiidae.
Pan- Pac.
Entomol.
34: 17- 32. 0013
Allred, Utah.
D. M. V.
1958b.
Mites found on mice of the genus Peromyscus in
Trombiculidae
and
miscellaneous
families.
17: 95- 102. 0014
Allred, sitol.
D. M.
47: 124.
1961.
Parasitic
mites
on
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55: 868- 71.
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Trombiculidae). J. M.
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New bat chiggers of the genus Perissopalla from
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J. M.
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0268
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J. M. 1969b. Five new lopadium ( Acarina: Trombiculidae).
Acarina:
0267
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Three new species of subgenus Parasecia Loomis
Brennan,
Venezuela
0266
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Colicus, species
Med.
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Entomol.
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new
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56: 807- 12.
Neotropical to
included
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(
des-
Acarina:
7: 271- 72.
1970c.
1942, ( Acarina:
A review of Neoschoengastia colombiae Boshell with descriptions of two new species misidentified Trombiculidae:
Colicus).
J.
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7: 489-
92. 0269
Brennan,
J. M. 1970d. A Acarina: Trombiculidae).
small
Bull.
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of
Calif.
chiggers Acad.
Sci.
from
Surinam
69: 32- 37.
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
22
0270
chiggers
63: 1694-
Am.
Soc.
Entomol.
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Trombiculidae).
Acarina:
(
Two new genera and a new species of Neotropical
1970e.
J. M.
Brennan,
96. 0271
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guyannensis,
0272
birds.
of
0273
and
the
57: 666- 67.
Anomalaspis
1971c.
J. M.
Brennan,
Pseudoschoengastia
(
South
of
1952,
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questionable
a second intranasal chigger
Trombiculidae),
Parasitol.
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Neoschoengastia
VI.
chiggers:
Endoparasitic
Acarina:
n. (
sp.
esorhina
Ann.
Trombiculidae).
(Acarina:
Brazil
Proechimys
rat,
spiny
64: 212- 15.
1971b.
J. M.
Brennan,
from
chiggers
new
northern
Am.
Soc.
Entomol.
Three
1971a.
J. M.
Brennan,
American
Acarina:
with
Entomol.
Med.
J.
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species
8: 602- 04. 0274
T.
Brennan,
descriptions
scutum
Revival
two
of
species
(
ignota
Acarina:
J.
Trombiculidae).
Acarina:
(
Vanidicus
of
the
of
species
new
Brennan
Jones,
and
Venezuela,
from
genus
Med.
1961, and
Entomol.
Med.
J.
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( Acarina:
taxa
related
on
notes
the
1973.
J. M.
Brennan,
Trombicula
new
9: 16- 18.
Entomol.
0276
in
eyes
with
chiggers
for
two
A new genus and two new species of Venezuelan
1972b.
J. M.
Brennan,
genus
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58: 1178- 81.
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new
Fauran,
rotundiscutata
Trombiculidae).
0275
Buclypeus,
1972a.
J. K.
Brennan,
10: 107- 09. 0277
Brennan,
0278
J. K. &
Brennan,
D
0279
Brennan,
J. K. &
0280
Brennan, the
J. M. &
Western
species
and
additional Trombi-
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of
Chiggers
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1977a.
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K. L.
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The
1960.
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Pacific
( Acarina:
Utah
15: 1- 26.
Basin Nat.
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H. T.
Six
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1956.
Beck.
E.
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1971.
Amerson.
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Guatemala ( Acarina:
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63: 554- 66. 0281
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J. M. &
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Brennan, chiggers
Goff.
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J. M. & (
Acari:
1977b.
M. L.
The Neotropical
records
other
Trombiculidae).
Acarina:
0282
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J.
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Parasitol.
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genus Hoffmannina:
Panama
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Venezuela
63: 908- 14.
Three
new
monotypic
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J.
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Med.
Entomol.
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Crotiscus
14: 541- 44. 0283
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J. M. &
M. L.
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1978b. northern
Two
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South America.
J.
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J. M. &
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Acarina:
Brennan,
Brennan,
J. M. &
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J. M. &
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J. M. &
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0293
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0294
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Soc.
Biol.
the
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Am.
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two
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52: 7- 16.
Pseudoschoengastia and four new
Acarina:
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Trombiculidae).
J.
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E. R.
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Chiggers
1961b.
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B. W. I.
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New genera and species of chig-
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Four new chiggers from Argentina
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50: 698- 702.
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58: 796- 800.
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0295
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0292
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45: 421- 29.
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A
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Trombiculidae).
Neotropical
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Trombiculidae)
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VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
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Two
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1972b.
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0301
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59:
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The Neotropical
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0303
0304
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0306
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P. H.
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New records of Surinam and certain French Guiana chig-
1972. (
Brennan,
Bull.
J. E. M. H.
the
with
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of
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The
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The
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Intercutestrix
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0311
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A new species of
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Trombiculidae).
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On the life histories of 3 species of
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(
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0352
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On the life histories of three species
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0353
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0354
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P. R.
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the
(
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Scrub mites from mammals and birds,
three
P. R. Hsu. 1958a. Neoschoengastia from
new
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Acta
Zool.
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Studies on the trombiculid larvae of China.
Acta
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H. T. &
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Acta
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Species of trombiculid mites and their 10: 404- 15.
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Acta
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VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
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0357
0358
H. T.
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of
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Acta
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(
Some
1962a.
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P. R.
&
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( Acarina:
Whartonia
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11: 549- 57.
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the
to
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Two new species of
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H. T. &
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Acta
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the
on
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Acomatacarus
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(
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H. T. &
Chen,
14: 355- 60.
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Zool.
Acta
the
of
age
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var.
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The
1962b.
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&
(
in
Chinese
with English summary) 0361
in
the
to
on
climbing
The behavior of the trombiculid larvae
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P. R.
H. T. &
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Province.
Kwangtung
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in
(
Chinese
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the
genus
0364
Tragardhula
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0365
H. T.,
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bicula
the
to
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the
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mites
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0367
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BULL. SOC. VECTOR ECOL.
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Techniques for testing the attach-
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Prague,
Nat.
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0429
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Three
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B. L.
J. P. a
A. L. ,
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Ewing
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Domrow,
R.
0475
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Malaya
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Domrow,
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Acarologia
The
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Innisfail.
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Their mite
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The
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0478
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29):
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Nymphs
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83: 227- 40.
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Audy (
Trom-
Acarina,
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0472
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
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en
de
barbarus Oubangui-
45: 1924- 25.
Exot.
Doc.
der
broyats
1952.
avec
cunninghami
bei
58: 66- 71.
Gaillard.
1950)
Herbstgrasmilbe
-
Zurich
inoculees
Mylomys Bull.
Herbstgrasmilbe
Tierheilk.
souris
Andre, sur
Chari ( A. E. F.) .
epidemic
Die
37: 587- 90.
LeGac & sur
( Marc
striatus,
0707 * Gispin,
Gouck. against
Parrish. of
Landw.
Ziege.
0706
of
18: 298- 300.
Aust.
0705
1962.
Gilbert.
mosquitoes,
Moule &
Aust.
identification
0704
the
5: 105- 06. I. H.
for
H. R.
treatments
Queensland.
0703
Gouck &
on
55: 451- 52.
Gilbert,
Gill,
Nat.
repellents
clothing 0702
Notes
S.,
as
Entomol.
0701
1960. Southwest.
J. E.
mites Neerl.
in
the
transmission
Indones.
Morb.
of
Trop.
2: 23- 35.
0708
Gispin, indica,
225- 30.
R.
1950b.
fam.
The
virus
Trombiculidae).
of
murine
Doc.
typhus
Neerl.
in
mites
Indones.
(
Schoengastia
Morb.
Trop.
2:
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
52
0709 * Gispin,
R. &
typhus
and
1951.
Warsa.
R.
infection
natural
rats
of
in
typhus
Java.
of
shop
3: 155-
Morb.
Indones.
The
II.
identity Trop.
ectoparasites;
Neerl.
Doc.
typhus.
murine
Endemic rat
and
62.
0710
M. A.
Goates, Brigham
0711
Goff,
Goff,
from
0713
Goff,
0714
Goff,
Goff,
1975.
M. L.
Midway
the
M. L.
Goff,
A
M. L.
new
A
Two
1977c.
Goff,
1977d.
M. L.
A
1978a.
0718
Goff, and
in
the
owiensis
new
Goff,
in
Acarina:
(
8: 456.
Trombiculidae)
( Acarina:
Hawaiian
the
Trombi-
Islands.
J.
with
Kohls
Trombi-
Trombiculidae)
Radfordiana
to
the
( Acari:
known
nine
from Papua New Guinea
1952 and Novotrombicula
( Acari:
1947
( Guntheria)
key
15: 21- 25.
Entomol.
rostrata Womersley
and
( Acari:
Guntheria
of
species of
R.
of
( Acari:
63: 915- 17.
Guinea,
Med.
Trombi-
14: 243- 46.
Entomol.
Guntheria
of
species
New J.
A new
Med.
J.
( Acarina:
14: 38- 41.
Schoengastia
of
Parasitol.
new
Guntheria
of
species
species
J.
Papua
Womersley
Trombiculidae) .
J.
64: 886- 90.
Parasitol.
0719
site.
12: 52- 54.
Entomol.
mammals.
subgenus.
redescriptions
Med.
new
Two
from
1978b.
M. L.
test
Trombiculidae)
Entomol.
Toritrombicula
of
plover
species
J.
from New Guinea
Trombiculidae) species
new
Two
from Papua New Guinea. 0717
golden
Nevada
Acarina,
(
Entomol.
species
the
Med.
J.
chigger
of
Med.
J.
new
chiggers
Islands.
species
at
3: 1- 12.
13: 735- 37. 1977b.
M. L.
of
rats
Ser.
Biol .
records
from New Guinea.
M. L.
kangaroo
Hawaiian
Pacific
the
Entomol.
on
Bull .
Islands.
1977a.
from
culidae)
0716
New
northwestern
culidae)
0715
Sci.
1971.
M. L.
culidae)
Med.
Univ.
Young
from the 0712
Mites
1963.
1979a.
M. L.
New Guinea,
with
The
genus
descriptions
Odontacarus ( Acari: of
four
new
Trombiculidae)
J.
species.
Med.
in
Entomol.
15: 143- 54. 0720
Goff,
M. L.
1979b.
New Guinea ( Acari:
0721
Goff,
M. L.
1979c.
Trombiculidae)
0722
Goff,
M. L.
culidae)
Entomol.
0723
Goff,
M. L.
Two
new
species
Trombiculidae).
Two
new
species
of
intranasal
J.
Med.
of
schoengastiine
from Papua New Guinea.
1979d.
infesting
A
new
murid
species
rodents
chiggers from Papua
Entomol.
J.
Papua
chiggers
Entomol.
Odontacarus
of
in
Med.
15: 533- 36.
New
(
Acari:
16: 112- 15.
( Acari:
Guinea.
TrombiJ.
Med.
16: 140- 41. 1979e.
Trombiculidae)
A
new
species
of
from Papua New Guinea.
Guntheria J.
Med.
( Guntheria)
Entomol.
( Acari:
16: 246- 47.
SEPTEMBER, 1956
0724
Goff, tion
J.
0725
BULL. SOC. VECTOR ECOL.
Notes
M. L.
Eltonella,
subgenus
( Eltonella)
with a redescripTrombiculidae). (Acari:
frittsi
16: 321- 24.
Entomol.
culidae)
the
on
Microtrombicula
of
Med.
Goff,
1979f.
M. L.
53
Host
1979g. infesting
Papua
New
1979h.
Species
of
by
exploitation
chiggers
land
Guinea
(
Acari:
Trombi-
Pac.
Insects
mammals.
20: 321- 53. 0726
Goff,
M. L.
horseshoe bat,
orange
( Acari:
chigger
Rhinonycteris
Trombiculidae) Rec.
aurantius.
West.
from the
Aust.
Mus.
8: 93- 96. 0727
Goff,
1979i.
K. L.
Acari:
A
new
and
genus
from
Trombiculidae)
Zyzomys
five
new
species
Rec.
argurus.
of
West.
chiggers
Aust.
Mus.
8: 81- 91.
0728
Goff,
from
1980a.
M. L. the
moss-
A
forest
new
species
of
chigger
Acari:
(
in Papua New Guinea.
rat
Int.
Trombiculidae)
Acarol.
J.
6: 85-
87. 0729
Goff,
1980b.
M. L.
species
0730
Goff,
Goff,
1980c.
M. L.
M. L.
from the J.
0732
Med.
Goff,
from
Goff,
white-
Goff,
The
A
Goff,
0736
Goff,
1981b.
Two
Med.
0737
Goff,
from
M. L.
Entomol.
Acari:
(
( Acari:
Trombicuidae)
Dasyuridae)
A
species
lowland
Entomol.
of
in Australia.
(
Entomol.
(
(
Acari:
Acari:
Trombi-
Trombiculidae)
18: 69- 72.
Guntheria
of
and
18: 33- 40.
Entomol.
melomys
penthetor
17: 494- 97.
chiggers
of
Trombiculidae)
Whartonia
chiggers
Med.
J.
of
Entomol.
Med.
J.
species
new
the
( Guntheria)
Rodentia:
Muridae)
( Acari:
in
New
18: 112- 15.
new
Colombia,
with
species a
key
Polylopadium ( Acari:
of
to
the
species
in
the
Trombi-
genus.
J.
18: 228- 31.
Entomol.
1981e.
descriptions
chigger
species
new
new
Two
1981d.
M. L.
Schoengastia
22: 35- 77.
Whartonia ( Acari:
of
Med.
J.
Islands.
from
Med.
J.
of
to
Ornithogastia
and
redescription
Five
1981c.
M. L.
culidae)
species
bats,
lectotype.
a
Trombiculidae)
key
21: 321- 27.
Insects
Pac.
species
a
Insects
Guntheria
from Papua New Guinea.
M. L.
Guinea.
new
fruit
1981a.
M. L.
Pac.
Trombi-
17: 477- 78.
Guinea of
rodents with
genera
new
Schoengastia ( Acari:
of
footed dunnart ( Marsupialia:
1980e.
M. L. New
A
from the Philippine
0735
species
Papua New Guinea.
Entomol.
culidae)
0734
in
1980d.
designation
0733
new
from New Guinea.
reported
Trombiculidae)
0731
Three
from Papua New Guinea
culidae)
of
The
three
18: 244- 48.
genus new
Teratothrix
species
and
a
(Acari:
key
to
Trombiculidae), the
species.
J.
with
Med.
BULL. SOC. VECTOR ECOL.
54
0738
Goff,
1981f.
M. L.
foliate
with
Goff,
new
species
from
setae
( Acari:
Guntheria
of
Papua
Guinea
New
Trombiculidae) Med.
J.
mammals.
18: 298- 99.
Entomol.
0739
A
body
VOL. 11, NO. 1
M. L.
Studies
1981g.
biculidae).
I.
Two
Papua
on
new
New
Guinea
chiggers
Schoengastia.
of
species
Acari:
(
J.
Med.
Trom-
Entomol.
18: 366- 69. 0740
Goff,
1981h.
M. L.
biculidae).
Studies
II.
Two
Papua
on
new
New
species
Guinea
chiggers
Guntheria.
of
Acari:
(
Med.
J.
Trom-
Entomol.
18: 370- 73. 0741
Goff,
1981i.
M. L.
biculidae). species bicula
0742
Goff,
from
0743
Goff,
in
the
K. L. a
0744
Goff,
0745
0746
Goff,
from
Incan "
of
chigger
(
notes
Acari:
Aust.
West.
Rec.
key
on
to the
Novotrom-
Trombiculidae)
8: 585- 88.
Mus.
Pseudoschoengastia ( Acari:
of
pocket
spiny Entomol.
mouse
(
Rodentia:
Trom-
Heteromyidae)
19: 12- 14.
Guinea and
and
of
ecology
545- 55,
Pp.
chiggers.
New
Guinea,
Vol.
in
J. L.
2.
Junk
new
species
opossum
(
Sasacarus ( Acari:
of
Marsupialia:
Trombiculidae)
Caenolestidae)
in Peru.
J.
Studies
IV.
on
Papua
Radfordiana of
a
new
New Guinea
is
species
a
junior
and
a
key
( Acari:
chiggers
synonym to
the
of
Trom-
Schoengastia
species.
Med.
J.
19: 147- 50.
M. L.
1982e.
biculidae).
Goff,
species
Guinea
Trom-
19: 60- 62.
description
Med.
A
rat"
1982d.
M. L.
Entomol.
0748
Acari:
(
18: 530- 35.
species
New
1982c.
M. L.
Entomol.
J.
chiggers
The Hague.
Med.
Goff,
new
Med.
J.
New
Australia.
Biogeography
ed. ,
biculidae).
0747
new
1982b.
M. L.
an
with
A
Papua
Entomol.
forest
a
Panama.
from
Goff,
A
New Guinea
new species of Guntheria with a
from
western
1982a.
K. L.
Gressitt, Publ. ,
in
Papua
on
Med.
J.
1981j.
skink
in
Two
genus
owiensis.
biculidae) taken
Studies
III.
V.
Studies
Entomol.
1982f.
M. L.
biculidae).
on
Papua New Guinea
A new species of Trombigastia
VI.
chiggers
(
Acari:
Trom-
from New Britain bats.
19: 204- 06. Studies A
new
on
Papua
species
New
Guinea
chiggers
Toritrombicula.
of
(
Acari:
Med.
J.
Trom-
Entomol.
19: 306- 08.
0749
Goff,
from
0750
Goff,
M. L. east
1982g.
M. L.
biculidae). 0751
Goff,
M. L.
biculidae)
Two
new
species
of
chiggers
African bats ( Chiroptera).
1982h. VII.
1982i.
Studies
on
Six
new
species.
new
species
A
from Venezuela.
J.
Papua
Med.
J.
Med.
New Guinea
J.
of
Med.
(
Acari:
Trombiculidae)
19: 376- 79.
Entomol.
chiggers
Entomol.
(
Paraschoengastia
Entomol.
Acari:
Trom-
19: 403- 12.
19: 556- 57.
( Acari:
Trom-
SEPTEMBER, 1986
0752
Goff,
BULL. SOC. VECTOR ECOL.
M. L.
1982j.
biculidae).
Studies
VII.
A
new
Papua
on
species
55
New Guinea
chiggers
Sasatrombicula.
of
J.
(
Acari:
Med.
Trom-
Entomol.
19: 601- 02.
0753
Goff,
M. L.
1983a. from
culidae)
Entomol.
0754
Goff,
0755
M. L.
1983b. from
M. L.
Goff,
Two
a
M. L.
new
shrew
1983c.
Trombiculidae)
0756
species
sheath-
Cotrombicula ( Acari: bat in Australia.
of
tailed
TrombiInt.
J.
species
Schoutedenichia ( Acari:
of
Insectivora)
(
in
Tanzania,
east
Africa.
TromInt.
25: 74- 78.
Entomol.
Goff,
new
common
25: 71- 73.
biculidae) J.
A
the
A
1983d.
biculidae).
new
species
from Java.
Int.
Studies
X.
Two
new
Entomol.
Papua
on
species
Gahrliepia
of
J.
New
( Gahrliepia)
25: 281- 84.
Guinea
chiggers
Schoengastiini.
of
( Acari:
J.
(
Acari:
Med.
Trom-
Entomol.
20: 40- 43.
0757
Goff,
from
1983e.
M. L. the
Entomol.
0758
Goff,
A
new
species
of
chigger
in
bat
sheath- tailed
common
(
Acari:
Trombiculidae)
Australia.
western
J.
Med.
20: 134- 35.
M. L.
1983f.
A
from
Trombiculidae)
new
species
Tanzania,
bat-
of
infesting
Africa.
east
J.
(
Acari:
Entomol.
20:
chigger
Med.
306- 08. 0759
Goff,
M. L.
0760
Goff,
M. L.
0761
Med.
Goff,
1983h.
0762
Goff,
from
M. L.
0763
Goff,
0764
Goff,
1983i.
A
A
1983j.
1983k.
M. L.
new
new
Two
M. L.
1984a. from
a
new
A
new
horseshoe
the
in
of
of
of
Apoloniinae
Doloisia
J.
of
Entomol.
species
bat
chiggers
Trom-
(
Acari:
Trom-
( Acari:
( Acari:
Med.
Trombiculidae)
Entomol.
20: 670- 72.
Austracarus ( Acari: Soc.
South.
Afr.
in
Korea.
Int.
Trombi-
46: 335- 40.
Sasatrombicula ( Acari:
collected
Trombi-
20: 618- 19.
Entomol.
Iran.
species
J.
( Acari:
20: 511- 13.
genus and subgenus Guntheria.
Med.
J.
species
Entomol.
New Guinea
of
species
taken
from South Africa.
culidae)
Papua
on
Zumptrombicula
of
Med.
J.
Africa.
migratory hamster
culidae)
species
A new species 20: 565- 67.
from Somalia,
M. L. a
Studies
XI.
Entomol.
culidae)
new
from South Africa.
biculidae). J.
A
1983g.
biculidae)
J.
Trombi-
Acarol.
10: 23- 25. 0765
Goff,
from 0766
Goff,
M. L. an
M. L.
Acari: Med.
1984b.
Egyptian
A
new
mole.
1984c.
species
J.
of
Gahrliepia
Entomol.
( Acari:
Trombiculidae)
21: 137- 39.
Three new species and new records of chiggers from the northwestern Hawaiian Islands. J.
Trombiculidae)
Entomol.
Med.
21: 257- 62.
0767
1984d.
M. L.
Goff,
biculidae).
Horwood
0769
Goff,
M. L.
Acari:
four
0771
Goff,
M. L.
Acari:
Goff,
K. L.
Acari:
0773
Goff,
0774
Goff,
M. L.
&
0776
Goff,
and
0777
Goff, Texas
0778
Goff,
0779
M. L. &
ina: Acad.
new
new
with
of
Trombi-
Beamerella
Mormoops
species
megalo-
species
of
Colicus
14: 565- 68.
Entomol.
a
genera
species
new
of
The
1982.
Odontacarus
of
pygosomal
Chatia Chatia
J.
plate.
and
Shunsennia,
( Acari:
a
Med.
J.
1983.
infesting
P.
of
Pseudoschoengastia
of
bird
A
new
and
Perissopalla
genus
three
of
description
beltrani.
Ingels.
J.
F. W.
tortoise,
M. L. &
Med.
Goff,
chiggers
Trombi-
new
precaria,
species
a
( Acari:
from
key
( Perissopalla)
Vene-
to the tuicali
19: 169- 75.
Entomol.
lizard
species
Eutrombicula
of
in Surinam.
J.
( Acari:
Med.
Ento-
20: 272- 74.
Acarina: J.
of
66: 835- 38.
the
synonymy
Trombiculidae) mol.
A
The
1980. a
of
descriptions
of
Hoffmanniella
K. L. &
bat,
Med.
J.
Venezuela
Brennan.
with
correction
species with
genus
species
new
Three
1978c.
from
Parasitol.
Trombiculidae), zuela,
1978b.
Brennan.
J. M.
&
from
14: 534- 35.
Entomol.
Brennan.
J. M.
J.
M. L.
Lipovsky
species
new
Venezuela ( Acari:
chinned
from Venezuela.
description
the
in
Ellis
15: 50- 51.
M. L. &
culidae).
Goff,
J.
A
leaf-
the
Med.
Trombiculidae)
2.
Speleocola
two
monotypic
from
1978a.
from
Brennan.
J. M.
Entomol.
with
0775
J. M.
&
Vol .
114- 19,
37: 501- 09.
Brennan.
Trombiculidae)
Acari: Med.
J. M.
in Colombia.
phylla,
0772
&
for
vectors
Pp.
VI,
genus of
A new
1977b.
Nat.
Trombiculidae)
The
Quadraseta
of
Basin
Great
culidae).
Trom-
Acari:
63: 1089- 91.
Brennan.
species
Guinea.
Acarology
eds. ,
descriptions
with
Parasitol.
new
(
trombiculid
of
New
Papua
1977a.
Brennan.
J. M.
H. L. &
Goff, and
J. M.
J.
chiggers
England.
Chichester,
&
in
Bowman,
Trombiculidae)
Venezuela.
0770
& C. E.
Ltd. ,
Guinea
15. 12 Distribution
rickettsiosis
Griffiths
New
21: 524- 26.
Entomol.
1984e.
chiggerborne
D. A.
Papua
on
A new species of Gahrliepia from the long- footed
XII.
M. L.
Goff,
Studies
Med.
J.
melomys.
0768
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
56
R. B.
Judd.
Loomis.
Trombiculidae)
Entomol.
N. L. &
R. B.
The first
berlandieri.
record of
Southwest.
a chigger from the Nat.
26: 83- 84.
1973. Two new species of Odontacarus Ewing from California and Baja California, Mexico.
10: 333- 36. Loomis.
Trombiculidae)
Sci.
1981.
Gopherus
from
73: 170- 71.
1974. central
A
new
United
species
States.
of
Comatacarus ( AcarBull.
South.
Calif.
SEPTEMBER, 1986
0780
BULL. SOC. VECTOR ECOL.
Goff,
M. L. &
R. B. Loomis. The genus Odontacarus ( Acari: 1977. from North America. J. Med. Entomol. 14: 370- 72.
biculidae)
0781
Goff,
M. L. & R. B.
Acari: North
0782
M. L.,
R. B.
Acarina: Goff,
J.
is
the
R. B.
Entomol.
J. M.
Loomis,
Med.
W. C.
1972.
synonym
Odontacarus dentatus
0.
of
Entomol.
1946)
Ewing,
galli
(
9: 479- 81.
Welbourn & W. J.
terminology ( Acari:
chigger
19: 603- 04.
Brennan.
senior
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0824
0825
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0826
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South
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(
in
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0870 * Hayashi,
S. &
in
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S.,
T.
Sanit.
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Hayashi,
S.,
T.
0874
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coastal
Med.
0875
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J.
Sanit.
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S.
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Geographical
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Teramura,
III.
Y.
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Studies
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An
a
(
of
the
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of
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Sasa.
Trombicula
the variations
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in Hok-
in Japanese)
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Queensland:
on
M.
studies on
larvae
unengorged
and Trombicula in Japanese)
43: 217- 22.
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1954. et
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Schluger,
the
of
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known
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2: 555- 64. 1941.
Med.
als
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Milben
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Igaku
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in Japanese)
Tanaka.
W. G.
fever
W. G.
tralia.
0876 * Hecht,
H.
Aust.
Heaslip,
Tanaka &
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Nisshin
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Japan.
scutellaris
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on
pomeranzevi
kaido.
(
Suzuki,
Bull.
Dep.
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al.
engorged
Trombicula
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Hayashi,
et
H.
with
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soil- surface
0873
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Studies
Health
Studies on the ecology of trombiculid
Studies on the surface distribution of Trom-
Suzuki,
of
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to
1954a.
1954.
II.
in Japanese with English summary)
distribution
0872
Suzuki.
scutellaris
93- 100. 0871
T.
Japan.
fever
in
Queensland,
north
Aus-
1: 380- 92.
Hautreaktionen auf die stiche blutsaugender Insekten allergische
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Zbl.
Haut-
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Om Leptus
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C.
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Herm.
Z.
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MacMillan Co. , G. W.
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0885
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snake
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in Japanese)
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0889
0890
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Bull.
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of
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(
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Y.
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acarine
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0896
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Hirst,
S.
S.
to
Myriapoda to
(
scorpions,
Mus.
Br.
man.
mites
(
Leptus,
Schoengastia,
17: 37- 39. domestic
animals
Br.
bees).
hive
of
and
injurious
parasitic
Hist.
injurious
disease
Mus.
(
( with
an
appendix
Hist. )
Nat.
Econ.
1- 107. The
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Hirst,
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On
1921.
S.
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and
Ser.
Econ.
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and
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"
10: 73- 77.
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S.
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the
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0892
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1915c.
S.
Hirst,
1926a.
Neotrombicula)
On
a
harvest
beings
at
bug (
Leeuwenhoekia
Sidney,
New
South
australiensis,
Wales.
Trans.
sp. R.
19: 150- 52. the
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S.
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0899
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0900
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1928.
Soc.
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On
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new
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the
on
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Soc.
some Australasian 5: 1021- 34.
1929a.
Sambon =
of
65
species
" tea
tree
Hatori
pseudoakamushi,
including
London
Trombidiidae.
of
itch
?).
mite"
Ann.
Hirst,
S.
that
1929b. the
causes
On
South Australia.
0902
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S.
London
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S.
1929d.
hirsti
Trans.
R.
L.
Hist.
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Soc.
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Hoeppli,
H. H.
to
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new
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north
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histochemical
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to
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" natural"
and
"
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13: 419- 28.
Neoschoengastia in
of
54- 58.
p.
special
fowls
of
Histological Ann.
mites.
Schumacher.
with
mite"
possible carrier 22: 451- 52. Hyg.
Shumacher,
Tropenmed.
man
a
Med.
Trop.
itch
" scrub
trombiculid
Found.
R. &
the
Trop.
R. &
Am.
On
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Soc.
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0906
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Trombicula hirsti
(
Proc.
12: 165- 76.
0904 * Hoeppli,
0905
R.
mite
new
Additional notes on Australian mites of the family
with
bicula
Inst.
itch"
Trans.
1929c.
Trombidiidae,
larval
the
" scrub
(
Mag.
3: 564- 65. 0901
four
3: 825- 41.
Mexico).
nunezi
Rev.
sp. ( Acar.
n.
Inst.
Salubr.
Trombidiidae Enferm.
Trop.
5: 221- 25.
0907
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0908
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0909
A.
Nac.
A.
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Cien.
A.
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nuevo
Dos
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1949.
Mexicanos.
genero
de
trombiculido
Mexicano.
An.
4: 451- 57.
Mex.
Trop.
nuevas de
Contribuciones Primera
trombiculidos Mexicanos.
9: 177- 89.
parte.
al
Rev.
conocimiento
Soc.
Mex.
de Hist.
los
trombi-
Nat.
10:
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0911
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0912
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A.
1950.
Mexicanos.
A.
1951a.
Mexicanos.
A.
Contribuciones 2a
parte.
parte.
conocimiento
al
al
City)
City)
City)
los
trombi-
de
los
trombi-
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conocimiento
Ciencia ( Mex.
de
10: 148- 53.
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Ciencia ( Mex.
Contribuciones 4a
al
Ciencia ( Mex.
Contribuciones 3a
1951b.
Mexicanos.
parte.
de
los
11: 97- 103.
trombi-
0913
Cien.
( Mex.
de
los
trombi( Mex.
Ciencia
parte).
conocimiento ( 9a
de
los
trombi-
Acta
parte).
Zool.
9):
(
Rev.
A.
Hoffmann,
parasitos
lla
de
los
trombiAn.
los
Esc.
trombi-
Folia
parte.
( Acarina).
p.
Ento-
Proc.
batracios
de
1st.
Int.
1024- 27. y
reptiles
en
11: 209- 16.
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Microbiol.
Lat.- Amer.
Estudio monografico de los trombiculido de Mexico
1970.
Trombiculidae)
Acarina:
de
parte).
conocimiento
al
New York.
Pr. ,
Acaros
1969.
A.
Mexico.
( l0a
Trombiculidae)
Mexican Trombiculidae Pergamon
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Hoffmann,
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3- 18.
1966.
A.
Hoffmann,
primera
An.
parte.
Nac.
Esc.
Biol.
Cien.
18: 191- 263.
Mex.
Folia
Trombiculidae).
Los
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Ang.
C. L.
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Hokari,
Cty.
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Singapore. Japanese)
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1945. Bull.
insects Ser.
Sci.
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The
1974.
C. L.
Hogue,
Ecological
1972.
A.
0924 * Hoffmann,
Mus.
Contribuciones
1965.
A.
al
Trombiculidae)
12: 101- 09.
Mex.
Biol.
Mexicanos ( Acarina:
Mex.
Congr.
0927
trombi-
Doctor Eduardo Caballero y Caballero.
Hom.
( Acarina:
Mexicanos
Hoffmann,
mol.
0926
al
Contribuciones
1963.
A.
culidos
0925
los
Ciencia
conocimiento ( 8a
Trombiculidae) .
Trombiculidae).
Hoffmann,
Nac.
0923
de
parte).
555- 58.
culidos
0922
7a
Una nueva especie Mexicana del genero Trombicula
1960c.
A.
Hoffmann, Acarina,
0921
Esc.
4: 1- 11.
Mex.
0920
al
Contribuciones
1960b.
A.
Mexicanos ( Acarina,
culidos
0919
trombi-
An.
20: 99- 105.
Hoffmann,
p.
trombi-
los
parte).
conocimiento (
Trombicul. )
Mexicanos ( Acarina,
City)
0918
Contribuciones
1960a.
A.
Hoffmann, culidos
0917
( 6a
los
16: 147- 48.
City) 0916
al
Trombicul.)
( Acar. ,
Mexicanos.
culidos
de
conocimiento
al
Contribuciones
1957.
A.
Hoffmann,
de
12: 87- 94.
City)
8: 17- 30.
Mex.
Biol.
conocimiento
( Mex.
Trombiculidae)
( Acarina,
Mexicanos.
Cien.
al
Ciencia
parte.
Contribuciones
1954.
A.
culidos
0915
5a
Mexicanos.
Hoffmann,
Nac.
Contribuciones
1952.
A.
Hoffmann, culidos
0914
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
66
(
Chicago
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Pr. ,
Los
of
the
27):
i-x
Costa
trombiculids
( Acarina:
23- 24: 116.
Mex.
Eutrombicula
Janzen,
on
observations
+
Angeles
Basin.
( Coloradillas, Natural
Rican
Chicago,
Nat.
Hist.
1- 173.
Pp.
chiggers) .
History,
Species
Ill.
Survey of Trombicula in Sime Road Internee Camp, Nampogun
Boekikyusui- Bu
Singapore
( 109):
1- 6.
(
in
loommirommom
SEPTEMBER, 1986
0928
BULL. SOC. VECTOR ECOL.
Holman,
L. J.
Acarina:
1973.
Continuous insemination in Eutrombicula splendens
Trombiculidae).
Proc.
eds. ,
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Int.
723- 25, Acarol . ,
Pp.
Congr.
in
M.
Daniel
Prague,
Prague.
0929
Horikawa,
0930
K.
1957.
in Mie
netsu"
Howard,
C. W.
1918.
Minnesota.
0931
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Study
Prefecture. A
H. T.
Chen.
0932
Hsu,
P. R. &
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Hsu,
Chen.
China
P. K. &
( 3):
H. T.
Kwangtung.
report
(
Rosicky,
Academia,
(
twenty- day)
in Japanese) the
on
Trombidiidae
of
17: 111- 44.
Sin.
9: 237- 54.
(
in Chinese with
The relationship between the tempera-
Trombicula
of
B.
On some species of GlobularoschoenZool.
1963.
development
and
Abstr.
0933
H. T.
Acta
1: 3- 7.
Minn.
1957.
Trombicula. English abstract) and
" Hatsuka
so- called
J.
preliminary
State Entomol.
Rep.
P. R. &
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the
on
Mie Med.
&
1971 .
akamushi
var.
deliensis.
Sci.
8- 9. Chen.
Acta
1964.
Entomol.
Eight new species of scrub mites from
Sin.
13: 737- 52.
(
in Chinese with English
summary) 0934
Hsu,
P. K.,
G. )
I. T.
Hsu &
Gahrliepia:
genus
G. (
agrariusa,
Study
T. R.
G. (
Ven.
1965. Five new species of the subhiangsiensis, G. ( G.) radiopunctata, G.
G.)
G.)
G. (
yunnanensis,
tsutsugamushi,
of
41).
part
G.)
Acta
megascutata from China. Zootaxon. Sin. 2: 74- 84.
in Chinese with English summary) 0935
Hsu,
P. R.,
of
rodent
I. T.
Hsu,
in
caves
Province.
Acta
C. C. the
Li & H. T.
" Shatien
Zool.
Sin.
Chen.
area"
of
16: 123- 31.
1964. Peal
(
The arthropod fauna
River
in
Delta,
Chinese
Kwangtung
with
English
summary) 0936
Hsu,
P. R.,
C. S.
Li & H. T.
Chen.
1963.
The effect of irradiation with
r- ray Co60 on the development and reproductive power of Trombicula akamushi
var.
deliensis.
Acta
Entomol.
Sin.
12: 578- 85.
(
in Chinese
with English summary) 0937
Hsu,
P. R.,
habits
T. C.
13: 56- 66. 0938
0939
Hsu,
(
P. R.,
K. C.
deliensis
Med.
J.
var.
K. C.
deliensis
Microbiol.
0940
Hsu, and
Y. C. its
Chinese)
Su & as
78: 276.
P. R.,
& H. T.
Chen.
akamushi
1964.
var.
Studies on the egg- laying
deliensis.
Acta
Entomol.
Sin.
in Chinese with English summary)
var.
Hsu,
Liu
Trombicula
of
(
Sin.
1954. relation
Chen. to
1959a.
the
Larvae of Trombicula akamushi
epidemiology
of
scrub
typhus.
Chin.
in Chinese)
Su & as
H. T.
related
H. T.
Chen.
related
7: 1- 9.
(
Studies to
to
1959b.
Larvae of Trombicula akamushi
the
epidemiology in Chinese) on
tsutsugamushi.
diseases.
Acta
Med.
of
Part
scrub
1.
Sin.
typhus.
Acta
The chigger mite 4: 260- 65. ( in
ctrl..
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
68
0941
Hsu,
3: 7- 28.
Sin.
0942
Hsu,
history
Hsu,
ification
mites
in
(
Studies
2.
Part
on
the Acta
Trombiculidae).
Acariformes:
(
species differentiation and class16: 101- 06.
Sin.
Zool.
Studies
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( Acariformes:
Acta
34.
Part
tsutsugamushi.
Microbiol.
Chinese)
The generic and
Trombiculindus
of
Acta
mites.
chigger
tsutsugamushi.
on
chigger
9: 452- 59.
1964.
Y. C.
of
Chinese)
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of
Sin.
Entomol.
0943
in
(
1959.
Y. C.
life
Classification
1957.
Y. C.
in
(
on
Chinese
with English summary) 0944
Hsu,
Trombiculidae,
Sin.
0945
Hsu,
Chekiang
Province
Walchia).
Acta
(
Trombiculidae,
family
in
(
chigger mite
Gahrliepia,
genus
6: 449- 60.
Sin.
Entomol.
Ento-
abstract)
A new species of
1956b.
Wen.
T. W.
Y. C. &
Acta
Acarina) .
order
sp. ( family 6: 129- 30. ( in Chinese with English
Acomatacarus mol.
Finding of adults and breeding place of
1956a.
Wen.
T. W.
Y. C. &
Chinese
from
subgen.
with
English
summary)
0946
Hsu,
deliensis Walch (
Med.
Chin.
0947
Hsu,
78: 282- 83.
J.
hoekinae ( Acariformes, Studies
Acariformes).
Chinese)
Revision
subfamily
the
of
and
Trombiculidae)
tsutsugamushi,
on
in
(
order
Trombiculidae,
family
1962.
Wen.
T. W.
Y. C. &
The life history and mating habits of
1959.
Wen.
T. W.
Y. C. &
Trombicula
description
of Sin.
33.
Acta
Entomol.
&Y. C.
Hsu.
1981.
part
Leeuwen-
a new genus.
12: 49- 53.
in Chinese with English summary) 0948
Hsu,
Wuyi
from Wuyishan
0949
Sci.
Huber,
0950
Huber,
Paris
of
of
A
new
species
of
Trombiculidae).
Reserve ( Acariformes:
in Chinese with English summary)
(
culture
index
an
as
jars.
to
relative
the
humidity
of
60: 289- 91.
Wash.
Soc.
Entomol.
Proc.
Prey attraction and immobilization by allomone from
1978.
I.
nymphs
Natural
Color
1958.
I.
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1: 214- 17.
J.
lie
H. X.
Liu,
S. L.
Y. H.,
Doloisia
Womersia
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(
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Acarologia
Trombiculidae).
20: 112- 15. 0951
Hubert,
lation
A. A. &
H. J.
1963a.
Baker.
Leptotrombidium
of
in
deliensis
Malaya
Studies
on
( Leptotrombidium)
the
Am.
Trombiculidae).
( Acarina:
habits
and L.
and
akamushi
J.
Hyg.
popu(
L.)
78:
131- 42. 0952
Hubert,
A. A. &
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Acarina:
0953
Hubert,
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17: 80- 81.
Baker.
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A. A., of
H. J.
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H. J.
Baker,
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Am.
C. C.
The
persistence
akamushi
J.
of
foci of Lepto-
transect
in
Malaya
78: 143- 49.
Hyg.
Chan &
disease
along
a
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the
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New data on the distribution of the mite Trom-
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Trombicula
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15(
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(
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1362
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L. C.
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1363
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1364
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A new
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1370
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A new
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Speleocola
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70: 102- 03.
Sci.
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species
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from
A
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Hot-
Mitsutomi,
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from
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mountain
S.
area
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mite,
Japan
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M.
the
II.
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Egashira,
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(
in
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J.
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Hist.
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55: 106- 19.
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Spring. with English summary) 1430
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( Walch).
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1428
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(
in
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1431 * Miura,
A.
biculid
1960a.
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proved
Studies
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their
life
the
on
by laboratory
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Japan.
result.
tsutsugamushi
of
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J.
( trom-
Descriptions
I.
Part
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Zool.
of
im-
11: 1- 5.
in Japanese with English summary) 1432 * Miura,
1433
1960b.
A.
biculid
Studies
by
mites)
in
life
the
on
laboratory
history
characters
11: 5- 14.
in Japanese with English summary)
* Miura,
( S. &
gamushi
M.
nymphs
1952.
Sasa.
and
( trom-
Descriptions
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adults.
structural
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of
Part
rearings.
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of
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by laboratory rearing.
J.
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3: 129.
(
in
Japanese)
1434 * Miura,
S. &
history
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M.
of
the
1960.
trombiculid
Further notes on the breeding and life Japan.
mites.
J.
Sanit.
Zool.
11: 59.
in Japanese)
1435 * Miyagawa, gamushi
Igakukai
B.,
I.
( river
Zasshi
Nagayo,
fever
of
168: 859.
D.
Mita
Japan) ,
ura
notes
& on
H.
Imamura.
the
mite
1916.
producing.
Tsutsu-
Taiwan
1436 * Miyagawa, Fukui.
1437
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Y.
Schimotuma.
Japan.
J.
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T.
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Miyajima,
M. &
fever),
the
Miyajima,
1440
M. &
Miyajima,
1441
1-
14.
(
M. &
in
(
On
on
rats
in
of
akamushi
in Japanese) ( Japanese
river
Trombidium
mite,
aka-
Japanese)
developmental
cycle
of
aka-
in Japanese)
(
life
the
fever.
river
(
the
of
The
1917a.
Japanese
T.
Okumura.
T.
K. &
a
Zasshi
cycle
Kitasato
Zasshi
Miyashima,
study 5- 38.
the
of
Arch.
" aka-
Med.
Exp.
Korea
Trombidium various
ihre
verwandte
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and
found
mites
allied
in
Japan.
in Japanese)
1917.
Trombidium
Formosa.
and
Studien ueber Japanand
893- 908.
the
(
Okumura.
Japan,
Akamushis
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( 254):
T.
K. &
from
Vergleichende
( 266):
Okumura.
comparative
Saikingaku
1917b.
Formosaische
and
Saikingaku
Miyashima,
mites
16.
1916c.
Okumura. of
Koreaische
Species.
1443
T.
development
Tsutsugamushi
( 253) : 1091- 106.
Zasshi
found
Japanese)
in Japanese)
Miyajima,
forms,
( 253) : 1-
Zasshi
mites
in
the
form
adult
Okumura.
Carrier
ische,
1442
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M. &
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1:
egg,
On
1916b.
and
the
(
1160: 1540- 41.
Jiho
Okumura.
Saikingaku
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5: 47- 48.
1916a.
Ikai
nymph
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1439
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On
1954.
Zool .
Okumura.
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preliminary
1438
VOL. 11, NO. 1
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akamushi
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371-
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1444
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1446
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C. F.,
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1448
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Mohr,
Mohr, sites
1450
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Chen,
C. O.
C. O.
C. O.
new
in
Med.
species
the
of
(
T. S.
Y. L.
Ho &
Discovery of the genus
Loochoo
Islands
( Acarina:
Trom-
10: 93- 99.
2: 251- 53.
trombiculid
mite
from
Tsinghai.
in Chinese with English summary) 1959.
Li.
Trombiculidae)
from
Three
new
China.
species
Acta
of
Entomol.
on
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rats
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Comparative
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small
New
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Am.
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Midl.
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1961. range.
Exp.
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1959.
from
standard
(
1956.
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chiggers,
( Acarina:
1947.
Luzon.
native
1449
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A Sin.
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Sin.
bat
Ewing,
biculidae).
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Relation J.
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of
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45: 154. load
to
host
size
and
SEPTEMBER, 1986
1451
Mohr,
BULL. SOC. VECTOR ECOL.
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1967.
areas
Acarina) .
1452
Mohr,
Am.
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tations
home
Conf.
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1964a.
Resour.
load
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and
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of
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as
29th.
Trans.
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181- 95.
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1964b.
Relation of
California
of
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Louse
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of
and
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home
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areas
Relation
small
Stumpf.
Nat.
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in
area
size
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to
Ecol.
Resour.
host
to
related
1454
Nat.
coarse-
Stumpf.
home
activity, topography, and parasitism in haired rats , and Trombiculidae
of
Guinea
Popul.
W. A.
and
Wildl.
Mohr,
N.
Res.
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New
of
105
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J.
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1455
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1456
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T.
1458
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of
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Exp.
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Taiwan
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Med.
Sotokufu
tsutsu-
Chuokenkyusho
Eisei-
disease
Pescadores.
of
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On
some to
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6th.
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Leg
Sci.
(
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on
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in
Formosa,
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of
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in
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Congr.
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Formosa
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"
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Assoc.
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1469
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Some
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1970.
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Russian
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Trombiculidae
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(
( Acari-
in Russian
with English summary) 1473
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in
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On
the
Minor.
( Acariformes,
trombiculids
Parazitologiya
Trombi-
6: 540- 45.
( Leningr.)
Russian)
Mulyarskaya,
formes,
(
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Three
from
species
new
Azerbaijan.
of
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L. V.,
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1476
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T. A.
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1972.
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of
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Kobystan.
Med.
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New
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4: 365- 70.
(
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Russian
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English
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1478
BULL. SOC. VECTOR ECOL.
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(
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in
1971. Nak-
the
Russian
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A. G.
Babaev,
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(
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1481 * Mulyarskaya,
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On
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Z. F.
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1971.
from
the
(
Russian)
in
Specific
Lencoransk
Small
composition
Zone.
Izv.
Cau-
small
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Nauk
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Province
1: 95- 101.
1965.
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L. V. ,
1967.
Azerbaijan
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Acariformes,
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SSR Biol.
the role of micromammals
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the
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A. S.
L. V.,
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-
(
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&
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Izv.
Malmedov.
Nauk
Zool .
Chirkova.
N. V.
Microtrombicula
genus
Azerbaijan.
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M.
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the
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Toritrombicula
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Southeast
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and
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Entomol.
1968a.
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with
five
of
from
9: 140- 51.
densipiliata
species
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Trombiculidae)
the
on
neotype,
Tori.
of
new
of
re-
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( Acarina:
Acarologia
region.
designation
with
Descriptions
Neoschoengastia
Australian
ental-
1492
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M.
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on
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the
a
genus
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species
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from
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TromAcaro-
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Nadchatram,
1968b.
M.
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A
in
for
technique
tropical
a
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Med.
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1495
1496
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1498
1500
1502
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Trombicu-
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Correlation
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J.
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1972.
M.
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Med.
J.
Ento-
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1979b.
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(
of
the
genus
Prostigmata:
and
Trombi-
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Gunong Benom Gunong Benom (
Hist.
from India M.
synonymic
The
from
Nat.
I.
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Entomol.
Mus.
Nadchatram,
( Zool. )
new
Expedition
1967.
Prostigmata;
9.
A collec-
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23: 189- 198.
species
Indonesia.
of
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Oriental
Ectoparasites
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mites
Insects
Malaysian
(
Acari:
Trom-
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Malay.
33: 75- 78.
Nadchatram,
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the
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1970c.
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Bull.
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M.
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An
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twelve
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1970a.
M.
of
M.
genera
Recent
1979c. of
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1505
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20: 174- 76.
15. 11 The natural history of the genus LeptoTrombiculidae)
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1507
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A
Bull.
key
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Southeast
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Inst.
Asian
Med.
to the
Res.
sub-
( Acari:
chiggers
Malaysia
( 16) :
1- 67. 1509
Nadchatram,
1510
Nadchatram, of
Pulau
M. &
M. &
1515
on
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of
Lauren-
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Acarina
of
the
mam-
129- 40.
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Entomol.
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new
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Entomol.
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Two
(
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The
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1916e.
On
Trom-
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(
in
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1542
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1612
1613
1614
1615
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1618
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6:
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M.
control
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of
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introduction
to
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Pr.
Tokyo
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classification,
p.
i-vi
+
1- 486.
(
in
Japanese)
1763 * Sasa,
K.
Japan.
1764
Sasa,
J.
K.,
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1973.
A
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K.
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24: 187- 95. Hayashi, on
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42.
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Kano &
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of
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trom-
2: 59- 64.
SEPTEMBER, 1986
1765
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1766
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of
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S.
Sasa,
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32.
the
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K.
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21.
part
R.
their
and
in
part
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Tokyo Med.
Hayashi,
Islands
3: 133- 34. (
1767
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tsutsugamushi,
on
On
Kano &
N.
(
from
1952.
the moun-
in Japanese) 1952.
Kumada.
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J.
(
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1768
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(
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1770
M.,
S.
M.,
Tokyo
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S.
Med.
M.,
Hayashi,
mites
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S.
M.,
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37.
I.
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species.
1776
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species
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1952.
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J.
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88.
J.
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A.
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the
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collected
1955.
in
on tsu-
Fuji
1950.
the
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(
Studies
( trombiculid
42: 439- 50.
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Mt.
Studies
Fuji
area.
(
1953.
to
the
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trombiculid
fauna
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1957a.
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1957b.
local J.
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25: 247- 54.
in
(
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1955. nymph
Fuji
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on
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8: 135- 40.
Miura.
larva
the
found
Studies
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Sanit.
Iimuro
Notes mites)
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the
biculid
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Tokyo Med.
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M. &
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S.
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tenjin
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Further
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in
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5.
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of
(
T.
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J.
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1774 * Sass,
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1773
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On
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S.
20.
part
in Japanese)
J.
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1772
6. (
tsugamushi,
1771
in
moles
Hayashi,
part
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Sasa,
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in Japanese)
gamushi,
J.
G.
tsutsugamushi,
from
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61.
Hayashi,
(
a
part
and
93.
related
in Japanese)
Studies of
area
in Japanese)
(
nagayoi
new
on
tsutsugamushi.
tsutsugamushi
Leeuwenhoekiinae).
( trom-
Japan.
1777
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97.
Sasa
et
1779
1950,
al.
Sasa,
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Acad.
R.
M. &
bicula
the
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in
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Trombicula
ecological
The
(
notes
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with
the
in Japanese)
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mites
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28: 247- 321.
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Studies
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discovered
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Exp.
Fourth
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Part
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in
(
Japanese)
M.,
1780 * Sasa,
R.
Asanuma.
and
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their
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1781 * Sasa,
in
Sasa,
M.,
1783
(
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Sasa,
M. &
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on
Health
Dep.
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On
1953. Kanagawa
Rep.
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the
H.
tsu-
Pref.
Studies
1951.
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M.
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N.
Annu.
A.
Kumada,
N.
Asanuma.
Y.
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in
(
Hyg.
on
tsutsu-
Med.
Tokyo
in Hokkaido.
68:
J.
(
in Japanese)
F.
Kawashima.
new
F.
1786 * Sass,
M.,
(
of
chiggers
Kochi
on
tsutsugamushi,
Tokyo
area.
Tokyo
J.
part
tsutsugamushi,
Prefecture.
part
Med.
Med.
15. 68:
J.
on
part
tsutsugamushi,
Tokyo
found in Kochi Prefecture.
M.
from
western
1952.
Egashira.
Trombicula
25.
sp.
Studies
17.
Med.
J.
Japanese)
Kawashima &
part
new
genus,
in
F.
gamushi,
from
on
Tokyo
the
Studies
1951b.
Kawashima.
species
68: 15- 17.
in
1951a.
chigger
of
birds
from
Studies
1952.
Obata.
Japanese)
M. &
Sasa,
T.
collected
species
new
Two
found
the
Japanese)
The
14.
Chiggers
9- 10. 1785
(
Kano &
R.
69: 557- 59.
A
On
in Japanese)
28.
1784
p.
Suzuki,
Tokunaga &
K.
in
Kano,
R.
K.
I.
shichito- netsu.
T.
Hayashi,
S.
Kanagawa
part
gamushi,
to
Sato &
Miura, Part
Japanese)
11
1952:
fever.
tsutsugamushi
the
of
A.
Kumada,
N.
shichito
on
relation
Miyagawa,
tsugamushi
Lab.
ecology
Kano,
R. T.
Tanaka,
9.
in
(
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S.
Studies
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p.
1782
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1952.
( Miyatrombicula)
Japan.
Tokyo
Med.
Studies
tsutsu-
on
kochiensis
new
69: 337- 38.
J.
sub-
(
in
Japanese)
1787
Sasa,
M.,
gamushi,
Kochi
1788 * Sasa,
F. part
Kawashima &
Tokyo Med.
Prefecture.
M.,
F.
Kawashima &
On
24.
gamushi,
part
epidemic
area
23- 24.
in Japanese)
(
of
S.
"
S.
J.
of
trombiculid
69: 43.
Hiromatsu.
the wild
Izumi"
1952a.
Hiromatsu.
A new species
18.
mice
and
(
Studies
mite
on
from
a
tsutsu-
mole
in
in Japanese)
1952b.
Studies
trombiculid
fever in Mie Prefecture.
on
mites
Tokyo Med.
tsutsu-
from
the
J.
69:
SEPTEMBER, 1986
1789
BULL. SOC. VECTOR ECOL.
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H.
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82- 97.
p. M. &
N.
Sasa,
Sasa,
M.,
Kumada,
N.
On
12.
K.
Sato,
S.
Shichito
fever,
on
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Island.
Hayashi,
Part
II.
T.
ZooHealth
Dep.
Japanese)
1952. in
Studies
forests
the
Hayashi,
S.
larval
tsutsugamushi,
on
Owu
of
Kumada &
in
(
A.
Y.
part
District.
Tokyo
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29.
On
Med.
J.
n.
sp.
Y.
Obata.
from
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1951.
Miura.
Studies
on
Tokyo
Med.
Euschoengastia.
genus
S.
Enomoto,
Trombicula
mite,
2: 1- 5.
the
Miura,
on
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new
Dobutsu
A.
Studies
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in
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in
N. A
Eisei
1792
(
M. ,
1951.
(
Kumada.
trombiculid
69: 661- 63.
1791
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1953.
Trombicula
of
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A.
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I.
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129
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part
68: 19.
J.
(
in
Japanese)
1793
Sasa, on
M. ,
Kumada,
N.
tsutsugamushi,
districts. 1794 * Sass,
M.,
Sasa,
Kumada,
N.
New
Tokyo Med.
1795
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Miura,
27.
Tokyo Med.
13.
part
A.
part
F.
Chiggers
69: 499.
J.
S.
Teramura.
species
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J.
(
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Japan.
1796
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J.
1797
Sasa,
A.
the
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A.
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Niigata
1955.
and
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Sasa,
nymphs
1953a.
Kinki
on
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tsutsugamushi,
Akita
on
the
prefectures.
tsutsugamushi,
Kawashima
Acarina,
(
Studies
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history of Japan. laboratory.
in
in
Japanese)
1951. in
Miura.
of
Miura.
the
on
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25: 159- 65.
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Exp.
M. &
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(
1952.
Miura.
Japanese)
Trombicula ( Miyatrombicula)
1952,
Suzuki & E.
T.
1953b.
Miura.
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on
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J.
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Exp.
Recent
34.
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1798 * Sasa, on
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A.
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(
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1956.
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Miura,
of
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H.
cluster
Japan.
A.
Med.
Exp.
of
Japan,
25: 197- 209.
Studies
on
(
83.
Notes
in Japanese)
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kuroshio.
Trombicula
part
and their laboratory
part
Japan.
J.
89.
Des-
Exp.
Med.
in Japanese)
A.
1921.
J.
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on
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Studies
1955.
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Japan.
rearings.
1799
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A.
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J.
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Tanaka &
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S.
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Tokyo Med.
Zool.
J.
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1955.
Ueno.
Y.
of
Trombicula
6: 135- 36. 1951. of
(
(
on
Nagayo
the
et
in Japanese)
Studies
Trombicula
in
Studies
scutellaris
Japanese)
on
tsutsugamushi,
akamushi
in
glass
1802
Sasa,
M. &
mites
Sasa,
&
1804
M. &
T.
1805
on
M.,
tsutsugamushi,
fecture.
1806 * Sasa,
1807 * Sass,
M. &
fauna
M.,
Sasa,
of
M.,
to
1809
1810
Sasa,
species
J.
68: 10.
Sasa, part
Y.
J.
Tokyo Med.
by
S.
2.
Chigger
cluster
mites
Gumma
Pre-
56.
part
The
Japan.
in
of
J.
Exp.
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on
especially
1950a. Island.
of
on the
to
expired
Studies
on
tsutsu-
unengorged with
and
reaction
of
refer-
to
carbon
27: 31- 43.
Med.
Studies
on
8.
part
from Japan.
Tokyo
larvae
special
tsutsugamushi,
unrecorded
Kano.
behavior
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akamushi,
J.
the
reaction
1957.
Hachijo
R. of
(
formation
Japan.
on
their
bionomics
Trombicula
1951.
Teramura &
and
Miura.
A.
the
hosts.
Notes Brumpt)
(
6: 166- 69.
Walchia
M.,
in
Honshu.
western
1955.
Zool.
of
Studies
1951.
collected
tsutsugamushi,
on
akamushi
previously in Japanese)
(
the
Japanese)
on
Teramura. of
On
Japanese)
formation
and
the
in
(
Prefecture,
Ueno &
mechanisms
S.
new
30.
part
Kumada.
N.
species
Teramura.
Notes
scutellaris
the
M. &
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Sanit.
J.
90.
expired
in
(
cluster
Tanaka,
H.
part
Trombicula
ence
Japan.
Peninsula.
Shimokita
new
Trombicula
of
their
Japan.
gamushi,
tsutsugamushi,
on
Studies
1953. Hiroshima
of
larvae
gas.
dioxide
breath.
of
102.
part
bird- parasitic
some
effect
Hayashi &
S.
68: 7- 8.
J.
Tanaka &
H.
mechanism
1808
the
the
Two
7.
4: 97- 100.
Zool.
from
Kano,
Takeoka.
E.
unengorged
CO2
R.
part
Tokyo Med.
trombiculid
Sanit.
of
in Japanese)
(
Savada,
T.
Studies
collected
mites
70: 143- 45.
Sasa,
1953.
Sato.
trombiculid
J.
by
trombiculid
tsutsugamushi,
on
appearance
rockhead
A new
53.
part
the
on
23: 333- 46.
Studies the
notes
with
28: 289- 91.
Med.
Sasa,
tsutsugamushi,
on
Med.
Exp.
on
the
on
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J.
1958.
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trombiculid
Exp.
Japan.
Osada.
Y.
Additional
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Miyatrombicula,
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M.
1953.
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K. of
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1803
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Med.
67: 22- 23.
J.
in Japanese)
1811
Sasa,
M.,
T.
gamushi,
Ueno,
together
with
Med.
1812 * Sasa,
K.
on
the
descriptions
Yeno,
1958. p.
H.
the
1957.
Tanaka.
bionomics
of
of
their
post-
Studies
on
Neoschoengastia
summer- tsutsugamushi
Tanaka,
Ecological
Studies
Islands. Tokyo.
Miura & H.
Notes
larval
of
Hachijo
tsutsuspp.
and
Islands, Japan.
J.
Suzuki &
H.
stages.
27: 217- 34.
M.,
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A.
hasagawai,
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96.
part
93- 101.
on
(
A.
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Miura,
fever.
in Japanese)
S.
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T.
on summer- tsutsugamushi General
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at
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SEPTEMBER, 1986
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Sasahara,
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1958a.
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Supplementary notes on the morphology of
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48: 247- 51.
(
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Japanese)
1814 * Sasahara,
T.
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Prefecture.
1958b.
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J.
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9: 73.
Kato.
1954.
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Sato,
A.,
K.
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Kyoto
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J.
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1816
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1959.
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1817
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1818
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Family Trombiin Pp. 118- 217, fauna
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30: 157- 64.
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Some
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cet
des
Schoengastia
important
Comp.
Revision
genre
(
Centra-
Acarina -
33: 619- 70. du
genre
Congo Belge Tervuren Ser.
Schoutedenichia
8.
65: 1- 101.
SEPTEMBER, 1986
2134
BULL. SOC. VECTOR ECOL.
Vercannen- Grandjean, tebolsi
n.
2135
Acarologia
Vercammen- Grandjean, rique
Acomatacarus
( Austracarus)
wit-
1: 253- 56. 1960a.
P. H.
septentrionale
sous- genres
1959.
un Trombiculidae larvaire parasite intradermicole de
sp. ,
la Taupe.
P. H.
155
nouveaux
et
(
Trombiculides Huit
centrale.
Acarina).
Arch.
larvaires
especes
Inst.
de
nouvelles
Pasteur
1' Af-
et
Alger.
six
38: 50-
69.
2136
Vercammen- Grandjean, larvaire
culidae
Acarologia
2137
manni
(
larvaire
P. H.
Schoutedenichia
de
Chlorotalpa
1960c.
sadini,
stuhlmanni
un Trombi( Acarina).
Acarina).
Gahrliepia
un
moucheti:
Trombi-
parasite des oreilles de Chlorotalpa stuhl-
nouveau,
Acarologia
Vercammen- Grandjean, une
1960b.
2: 210- 13.
Vercammen- Grandjean, culidae
2138
P. H.
parasite
P. H.
2: 214- 16.
1960d.
larve de Trombiculidae
Vatacarus ipoides Southcott 1957 et
apparentee
Eutrombicula.
a
Acarologia
2:
217- 23.
2139
Vercammen- Grandjean,
fication
Trombiculidae).
2140
Vercamen- Grandjean,
1952,
est
synonyme
1960e.
P. H.
Introduction a un essai de classi-
des larves de Trombiculinae Ewing 1944 ( Acarina Acarologia 2: 469- 71 + 1 unpaginated table.
rationelle
un
de
P. H.
Cheladonta Cheladonta
1960f. s.
Euschoengastia costulata Willmann et Cheladonta styriaca Kepka 1957 est
str.
costulata.
Bull.
Soc.
R.
Entomol.
nymphe
de
Multigniella
Belg.
96:
101- 07. 2141
Vercammen- Grandjean, tornis
Riv.
2142
P. H.
1960g.
Vercammen- Grandjean &
Parassitol.
La
Fain
1957
( Acarina,
cosme-
Trombiculidae).
21: 13- 20.
Vercammen- Grandjean, P. H. 1962a. Contribution a l ' etude de la faune d' Afganistan 75. I. Deux nouvelles especes de nymphes de Trombiculidae recueillies dans une grotte d' Afganistan. Acarologia 4:
362- 68.
2143
Vercammen- Grandjean, pour
Trombicula
l' importante
1962b.
P. H.
famille
A la recherche d' un neotype valable
Berlese
minor
des
le
1905,
Trombiculidae
representant
( Acarina).
disparu
Acarologia
de 4:
369- 80. 2144
Vercammen- Grandjean, Note
2145
preliminaire
Vercammen- Grandjean,
Heaslip 1943
est-
1963a.
P. H.
(Acarina:
il
P. H. une
Revision
Trombiculidae).
1963b. mythe?
(
du
genre
Trombigastia.
Acarologia
5: 57.
Le genre Myotrombicula Womersley et Trombiculidae -
Acarina).
Acarologia
5: 58- 60.
2146
Vercammen- Grandjean, nasal
249- 51.
du
lievre
P. H.
Africain
1963c.
Un
Schoutedenichia parasite intra-
( Trombiculidae -
Acarina).
Acarologia
5:
2147
1956
est
terrestris
( Trombiculidae
Schoutedenichia
veritable
un
Euschoengastia
1963d.
P. H.
Vercammen- Grandjean,
Gould
Acarina).
-
5: 394- 96.
Acarologia
2148
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
156
de
Trombiculidae
77 -
d' Afganistan
Contribution a l ' etude de la faune
1963e.
P. H.
Vercammen- Grandjean,
5: 582-
Acarologia
chiropteres.
615.
2149
including
Trombiculidae,
Pp.
Cornell
2150
J. A.
in
399- 407,
of
nymphs.
and
Vol.
Acarology,
Ascoschoengastia
1964a.
annulata
1.
Lawrence
Acarologia
( Trombiculidae).
Schoutedenichia
veritable
un
est
in
Advances
ed. ,
characters
larvae
New York.
P. H.
Vercammen- Grandjean,
1949
Naegele,
taxonomic
between
correlations
Ithaca,
Pr. ,
Univ.
Valuable
1963f.
P. H.
Vercammen- Grandjean,
6: 111- 13. 2151
Trombiculidae
2152
1964c.
P. H.
Vercammen- Grandjean,
1959
2154
d'
parasites
chiroptere
parasites
Jones
Brennan &
discors
Heaslip
( Trombi-
6: 127- 28. larvaires
Trombiculidae
Deux
6:
Acarologia
Guatemala ( Acarina).
du
Malaisie
6: 118- 22.
Womersley &
1964e.
P. H.
Vercammen- Grandjean, un
Trombicula
Acarologia
Acarina).
culidae -
de
Schoutedenichia
Deux
Myotrombicula
veritable
un
est
1964d.
P. H.
Vercammen- Grandjean,
Riedlinia
6: 114- 17.
Acarologia
Acarina).
d' Oiseaux ( Trombiculidae -
2153
nouveau
Acarologia
Acarina).
-
Un
1964b.
P. H.
Vercammen- Grandjean,
302- 08. 2155
sg. ,
n.
n.
6: 309- 11.
Acarologia
P. H.
Vercammen- Grandjean, Grandjean Revision
tibbettsi
un Trombiculidae larvaire parasite d' un chiroptere Coreen
sp.
Acarina).
2156
( Rudnicula)
Trombicula
1964f.
P. H.
Vercammen- Grandjean,
Brennan
&
du
genre
Le
1964g.
est
un
Riedlinia
genre de
synonyme
( Acarina
Trombigastia
Riedlinia
Vercammen
Oudemans
-
1914.
Acarologia
Trombiculidae).
6: 312- 23. 2157
Audy, fifty
1956, new
and
Microtrombicula
species
Leptotrombidium
Revision
1965a.
P. H.
Vercammen- Grandjean,
Ewing,
transferal
and
of
of
1950,
genera:
with
subgenus
Eltonella
descriptions
Chiroptella
Acarologia
Trombiculidae).
( Acarina,
the
to
of
genus
7( suppl.):
34- 257. 2158
Vercammen- Grandjean, an sp. , Acarina).
2159
interesting Acarologia
Vercammen- Grandjean,
of
commensal
7(
Tenotrombicula
1965b.
P. H.
suppl. ):
minteri
termites
n.
g. ,
n.
( Trombiculidae:
259- 65.
1965c.
P. H.
African
Iguanacarus,
a
new
subgenus
of
chigger mites from nasal fossae of the marine iguana in the Galapagos Islands,
with
Trombiculidae).
a
revision
of
Acarologia
the
7(
genus
suppl.):
Vatacarus
266- 74.
Southcott
( Acarina,
SEPTEMBER, 1986
2160
Vercammen- Grandjean, logia
2161
BULL. SOC. VECTOR ECOL.
7(
suppl.):
P. H.
in
Trois
1965e.
On
notes
Acaro-
rectificatives.
295- 96.
Vercammen- Grandjean, solenidion
1965d.
157
P. H.
chiggers
(
the
Trombiculidae:
true
nature
Acarina).
the
of
" striate"
Acarologia
7: 318-
20. 2162
Vercammen- Grandjean,
P. H. 1965f. Whartonia lepidopteriscuta n. sp. , noteworthy parasite of an African bat ( Acarina: Leeuwenhoekiidae). Acarologia 7( suppl.): 325- 28. a
2163
Vercammen- Grandjean,
extraordinary ina).
2164
Acarologia
1966a.
of
a
trombidiformes.
Vercammen- Grandjean,
1966b.
P. H.
Acarologia P. H.
pachywhartoni
n.
sp. ,
bat ( Leeuwenhoekiidae -
an
Acar-
Vercammen- Grandjean, Vercammen- Grandjean,
Notes
Note
Acarina).
1966d.
P. H.
de
terminologie
concernant
8: 285.
1966c.
trombidium ( Trombiculidae,
2166
Whartonia
Brazilian
8: 282- 84.
Vercammen- Grandjean,
les
2165
P. H.
parasite
a
propos
d' un
Acarologia
Revision of
1960 ( Trombiculidae -
nouveau
Erico-
8: 286.
the genus Herpetacarus
Acarina).
Acarologia
8:
631- 74. 2167
Vercammen- Grandjean, and
specificity Pp. 236- 39,
P. H.
Evolutionary problems of mite- host to
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Proc.
ed. ,
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on
Symposium
organisms.
Galapagos
Int.
Sci.
Berkeley. 1967a.
Notes
the
on
Trombiculidae.
Acaro-
9: 127- 34.
Vercammen- Grandjean,
Two Trombiculidae
of
viverrina
of
9: 135- 38.
P. H.
1947 ( Acarina:
Vercammen- Grandjean,
1967b.
P. H.
Acarologia
Vercammen- Grandjean, Hoffmann,
2171
Bowman,
Vercammen- Grandjean,
Transvaal.
2170
1966e.
relevance
Pr. ,
logia
2169
Univ.
R. L.
P. H.
Calif.
Proj . 2168
in
its
1967c.
Revision
Trombiculidae).
1968a.
P. H.
of
the genus
Acarologia
Riedlinia
Tecomatlana
9: 848- 64.
( Trombigastia)
libani,
a
new species indicating a relationship between the two subgenera Trombigastia 2172
and
Myotrombicula.
Vercammen- Grandjean,
Womersley
and
P. H.
Heaslip,
Ann.
Parasitol.
Hum.
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1943
Revision of ( Trombiculidae:
43: 59- 63.
Comp.
the genus Myotrombicula Acarina).
Acarologia
10: 65- 85. 2173
Vercammen- Grandjean,
Acarina: Dev.
2174
Command.
Wash. ,
Vercammen- Grandjean, phylogenie
Comp.
1968c.
P. H.
Trombiculidae &
et
de
44: 205- 10.
D. C.
P. H.
The
chigger mites
Leeuwenhoekiidae). p.
of
the
Army
Far East. Med.
Res.
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1969a.
taxonomie
U. S.
chez
Le
les
stade
larvaire,
acariens.
Ann.
sanctuaire de Parasitol.
la
Hum.
2175
Vercammen- Grandjean,
Etablissement
1969b.
P. H.
( Leptotrombidium)
Leptotrombidium
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akamushi
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from
chigger
Munich)
2178
a
sea
( 144):
1- 3.
snake
disparunguis ( Oudemans, Opusc.
2179
Zool. ( Munich)
Vercammen- Grandjean, Trombiculidae,
diformes).
2180
and
115):
1- 12.
ou
C. R.
Acad.
Sci.
( E.)
of
the
poppi,
a
Opusc.
Zool .
species
continents?
(
new
Walchia
Trombiculidae).
lignee de Trombi-
Acariens:
273: 2093- 96.
Paris
des
nepophylogenique
graphie
des
derive
Pilosite
1971d.
P. H.
Vercammen- Grandjean,
Int.
Budapest.
Kiado,
Sur la dispersion d' une
1971c.
P. H.
phoresie
tax-
2nd.
Proc.
group ( Acarina,
its
1929) (
pour ( Acarina:
phylogeny and
ed. ,
Akademiai
Revision
1971b.
P. H.
Vercammen- Grandjean,
lectotype 1910)
Trombiculidae) .
Acarina,
(
Evans,
Eutrombicula
1971a.
P. H.
Vercammen- Grandjean,
d' un
Brumpt,
Repository of
1969c.
P. H.
(
11: 94- 103.
Acarologia
Trombiculidae).
2176
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
158
appendiculaire
Acad.
C. R.
acariens.
et
chrono-
Paris
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273:
2267- 70. 2181
Trombiculidae)
Acarina,
2183
in
the
genus
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jamesoni
J.
Euschoengastia.
Prostigmata) ,
a
Angew.
58: 99- 103.
Zool.
P. H.
1971g.
Angew.
Zool.
Vercammen- Grandjean,
Z.
Vercammen- Grandjean,
1948 Med.
de neosomie
A
chigger
of
the
(Acarina,
Z.
subfamily.
zebra
Acarina,
(
Nouveaux regards sur les phenomenes
et de tachygenese chez
les
trombiculides
Bull.
Prostigmata).
Vatacarus ( Acarina:
africana
Apoloniinae
58: 173- 75.
1972a.
P. H.
the
of
representitive
new
Straelensia
1971f.
P. H.
Vercammen- Grandjean,
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2184
placed
8: 443- 45.
Entomol.
2182
Trombicula
1971e.
P. H.
Vercammen- Grandjean,
Acad.
du
sous- genre 58:
Sci.
Beige
R.
292- 94. 2185
Vercammen- Grandjean,
Parasitol.
2186
Folia
2187
P. H.
Parasitol.
P. H.
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Rosicky,
Proc.
demia,
Prague.
the
1972c. ( Prague)
1973a.
trombiculids.
nepophylogeny eds. ,
from
Asiatic- Pacific
region.
Folia
19: 227- 52.
( Prague)
Vercammen- Grandjean,
graphy.
Revision of Womersley' s Apoloniinae
1972b.
P. H.
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Acarina,
3rd.
Int.
Of
techniques
and
ortho- icono-
19: 289- 304.
Preliminary note in Pp. 311- 19,
Congr.
Acarol. ,
to M.
Prague,
a
tentative
Daniel
1971.
&
B.
Aca-
SEPTEMBER, 1986
2188
BULL. SOC. VECTOR ECOL.
Vercammen- Grandjean, Trombidiidae
1973b.
P. H.
Sur
( Acarina:
1815
Leach,
159
les
statuts
de
Prostigmata).
la
famille
des
Acarologia
15:
102- 114. 2189
Vercammen- Grandjean, eiplogue
Trombidioidea).
2190
Acarologia
1973d.
nudum
des
Neotrombidiidae
et
1905
Tragardh,
nilotica
15: 115- 20.
P. H.
nomen
Taxonomie
Blankaartia
probleme
Vercammen- Grandjean,
Cordisetoides
1973c.
P. H.
du
naturel
Morelacarus
Trombidiformes -
(
nomen
Acarina).
novum
for
Acarologia
15: 323. 2191
Vercammen- Grandjean,
P. H.
321- 28,
in M.
Pp.
graphy. Congr.
2192
Acarol. ,
1910),
Oudemans,
Vercammen- Grandjean, Ethiopian
2194
papilles genitales
2195
de "
Acarina).
2198
Andre
1954.
des
and
Med.
organs
de
1966a.
Andre.
M.
1966b.
Andre.
&
M.
Claparede
organs
Nouveau
of
the
et
les
respira-
Trombicula
ralphaudyi
Redescription
1966c.
Andre.
appliquee
n.
de
la
1910).
(Oudemans,
a
la
Introduction
systematique
des
a
la
Trombiculidae
8: 62- 70.
P. H. &
is
Andre.
M. true
a
et
Ann.
l'
P. H.
&
1967.
Cheladonta
J. R.
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P. H.
&
Audy.
( Acarina)
enlargissement
Parasitol.
Vercammen- Grandjean, XXIV.
Trombiculidae
89: 397- 439.
Afr.
Neoschoengastia
( Acarina,
brevi-
Trombiculidae).
9: 411- 14.
taxonomie
2200
1946,
Vercammen- Grandjean,
revision
P. H.
Acarologia
Acarologia
2199
16: 311- 18.
Eutrombicula ( Eutrombicula)
P. H. &
complexe"
Vercammen- Grandjean, palpus
M.
P. H. &
Vercammen- Grandjean, notion
larval
Bot.
verduni
Leeuwenhoekiidae
8: 55- 61.
Acarologia
2197
des
de Eutrombicula ( Eutrombicula) $ oeldii
adulte
Int.
17: 624- 30.
du Bresil
Vercammen- Grandjean,
forme
Leeuwenhoekia
Acarologia
Les
3rd.
8: 49- 54.
Acarologia
sp.
2196
originaire
de
famille
Some
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ortho- icono-
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la
1975b.
P. H.
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and
eds. ,
de certains acariens sont- ils des
Acarologia
toires.
Rosicky,
Prostigmata).
P. H.
Vercammen- Grandjean,
de
type
( Acari).
region
techniques
Revision
1974.
espece
1945 ( Acarina:
Womersley, 2193
P. H.
l'
Of
Academia,
1971.
Prague,
Vercammen- Grandjean,
1973e. Daniel & B.
du
Comp. J. R.
genre
1956. avec,
Note comme
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corollaire,
Schoutedenichia
Jad.
et
la la
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31: 427- 32.
Audy.
1957.
Malaysian
parasites
A revision of Heaslipia Ewing with descriptions of new species subspecies
Res.
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from
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153- 68.
(Acarina,
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Stud.
Inst.
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Vercammen- Grandjean, Eutrombicula
7(
2202
P. H. &
1965.
Audy.
Revision
of
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the
genus
Acarologia
280- 94.
suppl. ) :
Vercammen- Grandjean, culid
J. R.
( Acarina:
1938
Ewing
VOL. 11, NO. 1
mite
of
P. H.,
J. R.
undetermined
F.
Audy & and
genus
Trombi-
1966.
Pegazzano.
from Depok,
species
Java,
being
fragments in the Berlese collection misidentified as Trombicula minor
2203
Vercammen- Grandjean,
d'
parasite
2204
Acarologia
1905.
Berlese,
P. H. &
limace.
une
Vercammen- Grandjean, restrial
snail
a
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P. H.,
P. L. G.
host
new
for
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Un
1971.
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Benoit & J. J.
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trombiculid
aoutat
83: 180- 87.
Afr.
27:
Trop.
177.
2205
Vercammen- Grandjean,
from
Africa
east
Ann.
culidae).
2206
ina).
tres
et
Riv.
Zool.
de
deux
Bot.
Afr.
de
tion
d'
un
?
d'
et
2211
au
(
Acarologia
sa
new
Turkey.
des
Trombi-
un
genre.
Trombiculidae ( Acar-
Les
Fainiella;
Trombiculidae
J. B.
du
Revue
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-
&
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M.
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1966.
Acarologia
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1956.
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group
53: 343- 48.
brennani ( Jad.
Zool.
Descrip-
33: 5- 35.
Considerations sur un
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Jadin.
Trombiculidae
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genre,
J. B.
Hum.
1956a.
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Les
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genre
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1958.
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the
Description
of
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of
the
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1968.
Revision
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Acarina) .
10: 250- 68.
Vercammen- Grandjean, Two
(
de Gateria ( Giroudia)
Apoloniinae
subfamily
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Vercammen- Grandjean,
Multigniella:
1957b.
Fain.
&
P. H. &
Brunehaldia
species
famille
la
nouvelles.
Africaines
Vercammen- Grandjean,
new
2213
sujet
description de
subgenus
2212
P. H.
Vercammen- Grandjean, diverses
chiggers
56: 283- 92.
Myotrombicula.
Trombiculidae -
2210
A.
chiropteres.
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1957a.
Fain.
dans
especes
Vercammen- Grandjean, groupe
new
( Acarina:
18: 13- 16.
P. H. &
Vercammen- Grandjean, parasites
2209
heterogene
Eight
Trombigastia
50: 484- 96.
Am.
A.
1957.
Brennan.
genus,
Soc.
Parassitol.
description
2208
J. M. new
P. H. &
Vercammen- Grandjean,
Rev.
a
Entomol.
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2207
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2381
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H. &
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2382
S.
Aust.
A. K. &
notes
Woodward, mite-
on
in
2385
as
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2387
C. B.
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W. J.
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the
Soc.
S.
Aust.
71: 3- 12.
A snake den in Cedar Mountains Herpetologica
mites.
1946.
Tsutsugamushi
Philippine
1945.
12: 261- 68.
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scrub
or
Islands
during American reObservations on trombiculid mites.
II.
Vossbrinck.
arthropod
1977.
1974.
Notes
the
on
Scanning electron microscopy
Proc.
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Electron Microsc.
Biol.
Workshop
Appl.
SEM/-
2: 645- 651.
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Acarina:
(
Trombi-
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Euschoengastia.
Wrenn,
1956.
R.
32: 502- 13.
Scanning
Wrenn,
to
New genera and species of TrombiTrans.
parasitic
Philip. in
1944
T. A. &
tool
a
STEM.
2386
C. B.
Parasitol.
Woolley,
Parker. and
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D. D.
snakes
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borne
or
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2384
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2383
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W. J.
J.
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A
from the
culidae)
Kans.
new
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Entomol.
species
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of
beaver
in
47: 227- 38.
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Wrenn,
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2389
Wrenn,
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North
Wrenn,
W. J.
chiggers
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(
W. J. &
E.
Wrenn,
1976.
Two
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Loomis.
R. B.
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1967. Otorhinophila, a new genus of from western North America. Acaro-
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(
Acarina:
W. J. &
R. B.
C. E.
Bowman,
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(
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from
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eds. ,
North
The Euschoengastia radfordi species
Trombiculidae)
five
of
1973.
from
Ewing).
Eutrombicula ( Acari:
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Loomis.
genus
Trombiculidae)
Trombiculidae)
descriptions
2394
R. B.
the
Nat.
californica
complex
2393
R. B.
Acarina,
Acarina:
2392
&
in
9: 152- 78.
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mites
Southwest.
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Wrenn,
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2397
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(
in
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Japanese
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Part
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in Japanese)
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Distribution
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Y.
Part
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Shikoku
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Report
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1957.
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J.
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Japan.
M.
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1958.
Kagawa
Tsutsu-
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BULL. SOC. VECTOR ECOL.
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Shikoku.
Part
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3.
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2409 * Yamaguchi, culid
N.
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17S
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the
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Part
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Acta
Shikoku
Med.
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in Japanese with English summary) 2410 * Yamaguti,
S.
N.,
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gamushi
Inatoai
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Okayama
Kimura.
Districts.
1960. Japan.
Report
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the
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Zool.
11: 58.
in Japanese)
2411
Yamamoto,
S.
Miyazoki
1960.
Survey
Prefecture
on
chigger
Japan.
of
mites
Kagoshima
and
Med.
lice
in
12: 1-
J.
Hyuga,
6.
(
in
in
the
Japanese)
2412 * Yamamoto,
S.
Tojinwara Sanit.
2413
1961.
area
Zool.
Yamamoto,
12: 169- 73.
S. &
Kagoshima
An
K.
investigation
Kaseda
of
(
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mites
Prefecture.
Japan.
J.
in Japanese with English summary)
Kitahara.
Prefecture.
on
Kagoshima
City,
1964.
Japan.
Trombiculid survey in Ijuin area,
J.
Sanit.
Zool .
15: 119- 20.
(
in
Japanese)
2414
C. S.,
Yang, A
new
275.
2415
(
Yates,
2416
T. &
larval
Yu,
J.
2418 * Yu,
S. S.
Yu, of
Yu,
1959.
Trombicula kaohuensis sp.
from
mite
Chin.
Chekiang.
North
G. K.
Launchbaugh.
American
moles
(
1979.
Med.
nov. :
J.
78:
H.
Takahasi.
trombiculid
5: 61.
J.
1958.
mites
(
in
Chen &
Fuji
maneuver
Y. H.
Trombicula
animals
H.
Chou,
S. C.
deliensis
Sin.
7: 363- 72.
S. J. with
Lin.
Wu.
1957.
The Sin.
Lin & H. Y. and
(
Safety
the
Wu.
method
disease
in
Fukien.
Med.
survival and growth of mite 4: 245- 46. ( in Chinese) 1957. of
Study on the behavior
their
eradication.
Acta
in Chinese with English summary) 1957.
Rickettsia
in Fukien.
English summary)
Japan.
area.
in Japanese)
Acta Microbiol.
water.
S. S. &
J.
Seasonal occurrence and distribution
C.
infection
Talpidae).
16: 166- 68.
S. S.,
S. S.,
Ectoparasites from
Insectivora:
1957. Epidemiology of Trombicula 1: 22- 25. ( in Chinese)
Entomol.
2420
Wu.
Fukien
larvae in
2419
Pence &
of
Entomol.
Forces Med.
2417
K. H.
trombiculid
D. P.
species
Yatoai, of
Wu &
of
in Chinese) T. L.,
seven
Med.
C. L.
species
Study
on
tsutsugamuishi
Acta Microbiol.
Sin.
the condition of natural among mites and domestic 5: 425- 32.
(
in Chinese with
2421
Yu,
B. S. &
Entomol.
Yu,
between
two
9: 66- 74.
Sin.
Z. Z. &
1959.
Wu.
H. Y.
differences
2422
VOL. 11, NO. 1
BULL. SOC. VECTOR ECOL.
176
D.
(
and
morphological
on
deliensis
Trombicula
of
biological Acta
Walch.
in Chinese with English summary) A
1981.
Wang.
Q.
Yunnan ( Acarina:
Study
types
new
species
of
Neotrombicula
from
6: 273- 75.
Sin.
Acta Zootaxon.
Trombiculidae).
in
(
Chinese with English summary) 2423
2424
Yu,
Z. Z.
&
G. R.
Province
53- 56.
(
Yu,
Z. Z.,
G. R.
Zootaxon.
Yu,
Yu,
(
Conte).
2428
1958.
Yunker,
2431
2432
AscoActa
species
of
chigger
Trombiculidae) .
Three
new Acta
Acta
species
of
Zootaxon.
chigger Sin.
5:
Iowa
C. B. &
J. R. two
and
of
new
for
intranasal
chiggers
endothermal
laboratory
animals.
in
State Univ.
R. J. Pr. ,
1962.
species
(
Flynn,
Parasites
ed. ,
of
Ames.
Endoparasitic
Oudemans,
synoti
(Le
25: 31- 34.
technique
425- 92,
Brennan.
Doloisia
of
lucifugus
Myotis
of
47: 720.
Parasites
Animals.
mites
Wash.
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J. R.
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Med.
Yunker,
J.
Acarina:
chiggers:
II.
with descriptions of
1910,
a
Acaro-
Trombiculidae).
C. B.,
J. H. & J.
Four new chiggers ( Acarina: Trom1964. the epidemic area of Bolivian Hemorrhagic
1: 192- 95.
Entomol.
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1975.
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S. X.
B. B.
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two
new
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1961.
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host
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Chiroptera,
Acta
1980.
Soc.
Fever.
Zhao,
7:
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Tu.
The
1961.
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Trombiculidae).
2429
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China
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of
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D.
China ( Acarina:
C. B.
Yunker,
(
7: 270- 79.
G. R.
from
259- 63.
Yang & Z. Yunling
Sin.
Z. Z.,
mites
2427
Mt.
Chen.
Yunnan,
5: 150- 53.
G. R.
from
Zootaxon.
2426
new
Trombiculidae).
R. H.
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Sin.
Z. Z.,
mites
( Acarina:
in Chinese with English summary)
schoengastia
2425
Two
1982.
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Yunnan
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viral
Entomol.
and
(
chiggers,
Acarina:
Clifford,
rickettsial
12: 250- 55.
Endoparasitic
intranasal
species
C. M.
Philip,
of
chiggers:
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with descrip-
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J.
47: 995- 1000. 1982.
Entomol.
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A new species of Leptotrombidium parasitic on bats.
25: 102- 04.
(
in Chinese
with
English
summary)
SEPTEMBER, 1986
2435
BULL. SOC. VECTOR ECOL.
Zhao,
S.%.,
Walchia
Y.
24: 324- 26. 2436
Zhou,
(
M. ,
T. W.
3: 317- 19. Zhovtyi,
2438
(
the
N. D. 2.
2439 * Zhovtyi, On
p.
2440
I. F.,
Zumpt,
F. ,
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&
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Vost.
ed.
south
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1,
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O. A. (
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1956.
in
SSR.
Sin.
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1958.
Trombiculinae
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Emelyanova,
Izv.
of
collaboration
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n.
L. V.
Parazitol.
mites
Irktusk.
N. D.
I. F.
Dal'
the genus
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Pp.
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Sib.
East.
Konf.
the
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Acta
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Protivochumn.
for
Trans-
of
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2441
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I. F.,
logii.
1981.
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1 (
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Res. ,
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178- 184
BULL SOC. VECTOR ECOL., 11( 1):
EVIDENCE INCRIMINATING COQUILLETTIDIA PERTURBANS
DIPTERA: CULICIDAE) AS AN EPIZOOTIC VECTOR OF EASTERN EQUINE ENCEPHALITIS1
Isolation of EEE Virus From Cq. perturbans
I.
During an Epizootic Among Horses in New Jersey W. J. Crans2 and T. L. Schulze3
In 1983, four horse deaths due to Pastern Equine Encephalitis Virus( EEE) occurred on three
ABSTRACT:
farms in
separate
inland community
small
a
Ocean
of
County,
New
Jersey.
Mosquito trapping
revealed
that Coquillettidia perturbans was the dominant mosquito at each of the farms immediately after the equine deaths. FRE virus was isolated from host-seeking Cq. perturbans at two of the three farms where the equine deaths occurred during the preliminary survey period. Later survey revealed that FRP. virus was also present in Culiseta for
Cq.
collected
melanura
from the surrounding
12. 93 for Cs.
and
perturbans
area.
Minimum field infection rates of F.F.F.virus were 1. 63
Data suggest that Cs. melanura functioned as the enzootic
melanura.
vector during the outbreak period and that Cq. perturbans functioned as the primary vector to horses. August 26, and permitted an extended investigation of
INTRODUCTION
Pastern Equine Encephalitis( EEE) is recognized phenomenon in New Jersey primarily as a coastal with greatest virus activity in areas where freshwater swamps
border
salt marsh
wetlands (
1967, Goldfield and Sussman 1968). shown
that Culiseta
in bird
populations near
Altman
maintains
melanura
et al.
Research has F.F.F.
virus
swamp focus
the freshwater
the mosquito species present, the mosquito habitats in the area, and the incidence of EEE virus. Clark et al. (
1985)
gave a brief description of the outbreak
and the results of a study to test for transovarial transmission of FEE virus by Coquillettidia perturbans. This paper presents the results of the field
investigations that were initiated shortly after the first equine death was confirmed.
Morris et al. 1980, Crans et al. 1986) but that vectors
humans
to
and
horses
from
vary
one
MATERIALS AND METHODS
geographic area to the next ( Feemster et al. 1958, Chamberlain et al. 1969).
Aedes
The
salt marsh mosquito
has been incriminated as a primary FRF. in coastal areas of New Jersey ( Crans
sollicitans
vector of
1977, Crams common at
et al.
1986).
Sussman 1968, Sofield et horses have
Although
inland locations in the
equine cases are
state(
Goldfield
1983), the
al.
20
Four
mil)
area
to
of
the
of
occurred
cases
Jackson
Jackson Township is situated in the New Jersey pine barrens in northern Ocean County approximately 32 km ( 20 mi.) from the Atlantic coast. The area is characteristic of the outer coastal plain with sandy The soils composed primarily of Kirkwood Sand.
vegetation is typical of the New Jersey pine barrens,
been identified.
never
In 1983, five horse deaths due to FFF reported from the State of New Jersey( Schulze 1984).
and
vectors
Description of the Area
in
Township,
community in the northeastern portion The outbreak extended from County.
a
52
were
et al. km2
an
inland
of
Ocean
July
8
to
with considerable low shrub that is dominated by an oak and pine canopy. The area is drained by the Toms River and the South Branch of the Metedeconk River. At least 12 man-made lakes can be found in
the vicinity, most associated with cranberry farming. Aerial photographs and Geodetic survey maps show
New Jersey Agricultural Experiment Station Publication No. D-40101- 01-86 supported by U. S. Hatch Act with partial support from the New Jersey State Mosquito Control Commission.
2Mosquito Research and Control, Cook College, Rutgers University, P. O. Box 231, New Brunswick, New Jersey 08903, USA. 3New Jersey State
Department
of
Health, CN- 360, Room 705, Trenton, New
Jersey
08625, USA.
SEPTEMBER, 1986
more than where
BULL. SOC. VECTOR ECOL.
30 cranberry bogs in and around the were affected. A few of the larger
horses
systems
under
were
area
increase in
bog
Although the brain was taken at the time of death, inappropriate preservation did not allow confirmation
the time of the
cultivation at
179
titer to FFF
over
the
24 hr period.
survey, but the majority had been abandoned. One large tract of abandoned bogs was being utilized as a
by isolation of virus from brain tissue. The second
Fish
being raised on the same farm and had received
and
By
Game Wildlife Management Area. New
Jersey
standards,
Jackson
Township
is
quite rural with a population of
approximately 22, 000
people spread over an area of
260
The
is
region
noted
for raising horses
breed
of stables that
km2 (
100
mi
2).
with a number
for
and/ or train standardbreds
racing purposes. Zoning laws permit horses on land of limited acreage and many property owners keep one or more horses in a backyard situation. Horse
ownership is common even in developments the frontage may be less than 15 m( 50 ft.).
where
case involved a three month old Arabian colt that was
inactivated vaccine at the time of the first animal' s death. tissue.
The four in
of
fatalities
third
case occurred
in
an
aged,
development approximately 6. 4 km ( 4
mi.)
to the
Onset of symptoms were first noticed on August 12, and FEE was confirmed as the cause of
north.
death by isolation of virus from the brain. The final case occurred in a 16 month old standardbred that was
Equine Fatalities
equine
The
unvaccinated pony that was kept as a pet in a housing
being Confirmation
Onset occurred July 23, and the case was
confirmed by isolation of EEE virus from brain
trained at a special
facility.
The animal had
been vaccinated the previous week because of the F.F.F. virus
attributed to
publicity surrounding the prior cases in the area. The
encompassing the hamlets of Holmansville, Van Hiseville, and Jackson Mills ( Fig.
farm
1).
and 9.6 km( 6 mi.) from the first two cases. Onset of
occurred
Two
an
area
the animals were stabled in a
of
backyard farm. One the other was
for
trotters
available
being
and
kept in
was
raised at a
on the
being
raised
large training
four
two cases occurred on a 0.4
Holmansville.
development
TABLE
pacers.
information
a
on
1
ha ( 1. 0
and
facility
summarizes
The first
animals.
acre)
farm in
for
show.
The first
14
animal, a
CNS
month
symptoms
8. The attending veterinarian collected serum that date and again the following day just prior to
July
euthanization.
symptoms began August 26, when the owner noticed that the animal was less active than its stable mates. The horse was allowed to remain in the exercise area
overnight and was found dead the following morning. The case was reported immediately and confirmed by isolation of F.F.F.virus from the brain.
Both involved Arabians that were
old unvaccinated stallion, exhibited on
small
was located 3. 2 km ( 2 mi.) from the housing development where the pony contracted the infection
The blood
samples showed a
64- fold
Mosquito Collections
Mosquito surveillance was initiated as soon as possible at the farms where equine deaths were reported. Each of the sites was visited at least twice to
ascertain
the
mosquito
species
TABLE 1. Equine deaths due to FEE virus in Jackson Township, New Jersey during 1983. Date
Vaccination
of
Onset
7/ 8
7/ 23
8/ 12 +
8/ 26
Age 14
3
10
16
mo
mo
yr
mo
Breed&
Sex
Location
History
Arabian
Farm 1
Unvaccinated
Stallion
Hohnansville
Arabian
Farm 1
Colt
Holmansville
Grade Pony Gelding
Van Hiseville
Standardbred Stallion
Farm 2
Farm 3 Jackson Mills
Vaccinated Previous Week
Unvaccinated
Vaccinated Previous Week
present
once
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
180
At4P*
1
4
4
t iolibA, 11
SIP t
1\
lipilp
4'
P
Mors•
Farm 3
r
c.
Jackson
Hors•
Mills
VIP'
Farm 2
526
1111111V. S]
7
Vanhis• vill. Hors•
Farm 1
N• lmansvill• , r
525
r
2 mi
Figure 1. The Jackson Township area of Ocean County, New Jersey where Eastern Equine Encephalitis reached epizootic proportions in 1983.
confirmation of
of
FF.F. had been determined as the cause
death. Portable CDC traps baited
used
to
make
distribution
at
the
where
sites
horses generally
for
ice
were
were used to selectively collect Culiseta melanura over
wider
a
geographic
area
for
isolation
deaths had
were also used to
township to locate potential resting places of Cs.
equine
light traps
dry
attempts. The boxes were, at first, randomly scattered in damp forest and swamp habitat throughout the
collections
collections
Trapping
attempts.
the
farms
Jersey
the
supplement
where
the
New
occurred.
of
all
with
for
virus
included the
species
isolation
stable
area
spent the night, exercise
melanura.
When concentrations of the species were
located, the resting boxes were grouped in those areas that yielded the largest collections.
areas where the horses were kept during the day, and wooded
areas
September (
In early surrounding each farm. the last equine case) resting boxes
after
Virus Isolation Protocol All
of
the
mosquitoes collected
by
light
traps
SEPTEMBER, 1986
and
BULL. SOC. VECTOR ECOL.
resting boxes
laboratory species,
frozen
were
The
collection.
the
stage
New
laboratories chicks
blood
of
they
were
than
12
hours
Chamberlain
by
suspensions
State
Jersey
from
chicks
at the time of
by
date,
site,
digestion.
meal
17. Culiseta melanura comprised a somewhat larger percentage of the mosquito population at this and
in the
speciated
57° C(- 70° F)
where
less
described
ice
dry
were
on a chill table and pooled
and
specimens were stored at-
to
on
specimens
site due, primarily, to a local breeding source within
The
45
by
et al. (
the
1954).
yds)
the
of
August
30
stable
Farm 3 was
area.
September
and
1,
when Cq. perturbans were well into their decline. Anopheline
species formed a large portion of the collections at that site, but the total number of specimens trapped was low due to cool night time temperatures at that
methods
The brain
symptoms
showing
50
m (
sampled
and submitted
Department of Health tested in White Leghorn old
181
were
time of year.
innoculated into fresh chicks, suckling mice, and duck The final virus identification fluorescent by antibody ( FA) from the
embryo tissue culture.
Virus
was
Trap Collections A total of 1, 895 specimens were grouped in 67
made
tissue culture preparations.
pools
RESULTS
A
total of 1,
summarizes the
the on
fauns
farms.
light the
of
were
for the
trap
from
Culex
Cq.
death
equine
gorged
was a minor
Farm 1 (
apparently
season.
an
collections
locations.
was sampled
perturbans
by
early
season
that time of
but
July 20- 28, near
was
Culex
salinarius
Farm 2,
when
the
combined.
Two of
peak
salinarius
when
area
TABLE 2.
the
also was
major
sampled on
perturbans
representing approximately
total collection for that species on that trapping date. The minimum field infection rate ( MFIR) for Cq.
the
during this epizootic period was 1. 63
perturbans
isolations per 1, 000, or 1 isolation for each 612
were probably multi-parous Coquillettidia perturbans and
were
Cq.
pool contained eight unengorged Cq. perturbans, the
pop-
was
The first isolation was
perturbans.
The second isolation was obtained from Farm 3, 11 days after the equine death at that site. The positive
the site of
and
year.
TABLE 3 lists
attempts.
farms
Light
one- third of the mosquitoes collected on that date.
only other mosquito species present in large numbers. Small numbers of Aedes canadensis remained from the
isolation
the
obtained from the collection made at Farm 1, two days after the second horse died from EEE at that location. The positive pool contained 100 unen-
mosquito at each of
melanura
Cq.
unengorged
TABLE 2
shows that
results and
dominant
Culiseta
the
at all
2 equine deaths)
ulations
three
immediately following
in
species
by
specimens was collected
the
the
the premises.
present
Cq.
068
trapping
was
virus
From
the pools from the horse farms yielded EEE virus. Both of the isolations were obtained from pools of
CDC traps from
perturbans
for
Attempts
the results from the three
Mosquito Collections the
Isolation
tested.
Results of the trapping studies showed that Cq. perturbans was not only the specimens
species at
dominant mosquito species in the area, but that F.F.F
August 15
virus was present in host-seeking Cq. perturbans at
Mosquito collections from Jackson Township farms where equine deaths were investigated in 1983.
No.
Cq.
perturbans
613
(
Cx.
salinarius
107
(
Ae.
canadensis
27
Farm 3
Farm 2
Farm 1
Mosquito Species
(%)
No.
(%)
77)
147
(
13)
45
(
3)
No.
(%)
66)
22
( 20)
4
(
8)
0
(
0)
0
(
0)
( 44)
Cx.
restuans
18
(
2)
12
(
5)
0
(
0)
Ae.
vexans
18
(
2)
0
(
0)
0
(
0)
Cs.
melanura
4
(<
1)
13
Ae. triseriatus
4
(<
1)
2
An. bradleyi
3
(<
1)
Other*
1 (<
1)
An. punctipennis, Ae. trivittatus,
and
(
6)
4
(<
1)
1
0
(
0)
17
4
(
2)
2
Cx. pipiens.
(
8) 2)
(
34)
( (
4)
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
182
two
farms
three
the
of
from mosquitoes in that region in advance of the first
deaths had
equine
where
from the farms but the
collected
horse infection
numbers tested were
however,
very low.
Subsequent collections,
of the year.
failed to yield EEE virus from
the large
samples of Cq. perturbans that were tested. Francy Culiseta
From
Isolation
Virus
melanura
(
During
the
survey
Cs.
232
period,
from resting boxes were TABLE 4 lists the collections
period and shows that F.EE virus was
Cs.
melanura population
the
latter
the
part of
in Jackson
by
sampling present in the
Township during
the collections were
season when
The MFIR for the
made.
and a single
melanura
tested for FRR
collected virus.
investigated
1982)
an
outbreak
period
collection
was
12. 93, an exceptionally high rating for this species in New Jersey ( Veazey et al. 1980). Data, however, were only collected over a short period of time and the figure is probably not truly indicative of the amount of virus in this species during the outbreak period.
human
case.
was the dominant mosquito in that area and the
results of field investigations and virus isolation attempts
suggested
probable
that
Cq.
in
the
also
was
during EEE three
only
virus was
farms
Coquillettidia
dates.
species
mosquito
the time that
isolation peak
these
with
horses
found
positive
contracting EEE.
were
isolated from the
perturbans
EEE
virus
was
extremely low.
numbers
obtained when
when
for the
season
the species was
suggested that exceptionally high populations of Cq. perturbans might be necessary to initiate epizootic transmission of RF . virus by this species. Sofield et 1983) conducted field investigations at inland
al. (
sites where equine cases of EEE were confirmed in season.
was
Their results
the
dominant
mosquito at the farms where the equine deaths occurred but no virus was isolated from any of the specimens that were captured for virus isolation attempts.
Current knowledge
points to
Cs.
melanura
as
the primary maintenance and amplification vector of
FEE virus in any given area ( Burbutis and Jobbins 1957, al.
Kissling 1958, Hayes et al. 1962, Morris et
1980, Nasci
Edman 1981).
and
No virus was
detected in Cs. melanura collected at the farms during
the outbreak period but follow-up survey did show
near
and
the second was
that Cs. melanura were positive for EEE in that area and that the MFIR in Cs. melanura was extremely
well
into its
high.
species
was
seasonal
From the information available, it would appear
decline. Coquillettidia vector of
The information
One
the
obtained
the
species at two of the
where equine cases were contracted.
was
was
Cq. perturbans.was
extremely abundant in that area and the MFIR for
in New
involvement closely
activity period for Cq. Jersey. The advent of equine Jackson Township corresponded
as the peak
perturbans
However,
vector.
Jersey during the 1982 showed that Cq. perturbans
Data from Jackson Township strongly incriminate Cq. perturbans as the primary vector to horses during the outbreak period. Hagmann ( 1953) cites
June 23- July 5,
southwestern
Coquillettidia perturbans
New DISCUSSION
perturbans
in
Michigan during 1980 that involved numerous horses
Collected After the Outbreak
has been
perturbans
F.FF, since Howitt
suspected as a
et al. ( 1949)
that Cs. melanura
was functioning as the enzootic
isolated the
vector during the outbreak although the data only
in Georgia. Since
document enzootic involvement after the equine
that time, however,
deaths actually occurred. Data also suggest that Cq.
been
only limited field evidence has to directly link the species with either
perturbans was the primary vector to horses at the
virus
from
wild-caught specimens
gathered
equine or
human cases
Srihongse ( 1978) perturbans
the
in
fact that
of the
isolated
central
New York
this was the
TABLE
Morris
and
from
Cq.
inland focus during 1983. Sofield et al. ( 1983) point to the problems associated with sampling for EEE
and made mention of
virus after horse cases are confirmed. The extended
disease.
FRF
virus
first isolation
ever obtained
outbreak period in the Jackson Township area overted
3. Virus isolations from mosquitoes collected from three farms where equine deaths were attributed to FF.F.virus in 1983.
Mosquito Species
Cq. perturbans Other
Total
No.
EEE
Tested
Pools
Positive
MFIR
1, 225
15
2
1. 63
670
52
0
0
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
TABLE
4.
Virus isolations from Cs. melanura collected from Jackson Township, New Jersey following an epizootic in horses in 1983. Collection
No.
Period
Tested
September 08- 09 September 12- 15 September 19- 20 September 28- 29 October 10- 12 Totals aMFIR=
those problems and permitted equine case
in one instance
the others that
183
and
No.
FF.F.
Pools
Positive
18
3
67
8
1
81
5
2
47
4
0
12
2
4
232
22
0
3a
12.93
sampling prior to an immediate response to
followed.
Chamberlain, R. W., W. D. Sudia, P. H. Coleman, J.
G.
Johnson,
T.
and
H.
Work.
1969.
Arbovirus isolations from mosquitoes collected
in Waycross, Georgia, 1963, during an outbreak
Acknowledgements
of
eastern
equine
Am.
encephalitis.
J.
Epidemiol. 89: 82-88.
The authors thank Dr. Ronald Altman, Dr. William Parkin, Dr. Bernard Taylor, and Mr. Wayne Pizzuti of the New Jersey State Department of Health for their
Special
efforts and advice
appreciation
is
Mr. Walter Gusciora,
during
this
and
investigation.
Mr. David Adam,
extended to
Mr. Jeff Winegar
of the
New Jersey State Health Department for data that was pertinent to the of the
Ocean
fmdings
County
was most valuable,
of this study. The assistance Mosquito Control Commission
particularly in
initial location
the
of mosquito breeding habitats in the Jackson Township area. The study would not have been possible without the
on equine cases that were
Dr.
Sidney Nusbaum and Dr. Robert of the New Jersey Department of Agriculture.
supplied
Eisner
by
data
REFERENCES CITED Altman, R.
M., M. Goldfield, and O. Sussman. 1967. The impact of vector-borne viral diseases in the Middle Atlantic states. Med. Clin. North Amer. 51( 3): 661- 671.
Burbutis, P. P. melanura
in New
Kissling. western
and
C. L. Crabbs.
1985.
Absence of eastern equine encephalitis ( EEE) virus in immature Coquillettidia perturbans associated with equine cases of
F.F.F.. J. Am.
Mosq. Contr. Assoc. 1( 4): 540-542. Crans, W. J.
1977.
The status of Aedes sollicitans
as an epidemic vector of eastern equine encepha-
litis in New Jersey. Mosq. News 37( 1): 85- 89. Crans, W. J., J. McNelly, T. L. Schulze, and A. Main.
1986.
Isolation of eastern equine
encephalitis virus from Aedes sollicitans during
epizootic in southern New Jersey. Mosq. Contr. Assoc. 2( 1): 68- 72. an
J. Am.
Feemster, R. F., R. E. Wheeler, J. B. Daniels, H. D. Rose,
R. E.
Kissling,
R.
O. Hayes,
R. E.
Alexander, and W. A. Murray. 1958. Field and laboratory studies on equine encephalitis. New England J. Med. 259: 107- 113.
and
D. M. Jobbins.
1957.
Culiseta
and eastern equine encephalomyelitis
Jersey.
Proc. N. J.
Mosq.
Exterm.
Assoc. 44: 68- 70.
Chamberlain,
Clark, G. G., W. J. Crans,
R.
W.,
1954. equine
R.
Use
K.
Sikes,
of chicks
encephalitis
Immunol. 73: 106- 114.
in
and
R.
E.
Francy,
eastern and
and
J.
Sci.
studies.
D. B.
1982.
Eastern equine encepha-
lomeyelitis ( EEE) and consideration of some alternatives for dealing with EEE as a public health problem. Pp. 15- 18 ha: J. G. Engemann Ed.) Eastern equine encepha- lomyelitis ( F.F.F,) public
for
health in
southwestern
Citizens Ctr.
Mich. Univ., 26 pp.
of
SW
Michigan.
Mich., West.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
184
Goldfield, M.
outbreak of eastern encephalitis
Introduction
I.
perturbans
(
L.
L.,
R.
Sussman, logical
and
Biology Proc.
Walker).
Mansonia
of
N. J.
Mosq.
O.
Hess,
D.
1962. Entomo-
M. J. Bonese.
the 1959 outbreak of eastern
aspects of
encephalitis
A.
Beadle,
D.
in New
Jersey.
Am. J.
Trop.
Med.
Hyg. 11: 115- 121.
lomyelitis virus in up-state New York, USA.
relation
to distance from
Recovery
Bishop,
and
R. H.
of the virus of eastern
Gorrie.
1949.
equine
encephalomeyelitis perturbans)
from
mosquitoes
collected
in Georgia.
breeding
eastern Massachusetts eastern equine encephalitis
enzootic foci. J. Med. Entomol. 18( 6): 493- 500. W. J. Crans, and W. E. Parkin.
1984. Multi-agency response to eastern equine encephalitis
Jackson
in
Township,
R. E.
1958.
arthropod-borne
Host relationship
the
Ann. N. Y.
encephalitides.
Ocean
County, New Jersey. Proc. N. J. Mosq. Contr. Assoc. 71: 87- 93. Sofield, R. K., J.
Kissling,
J. Med.
Nasci, R. S. and J. D. Edman. 1981. Blood-feeding patterns of Culiseta melanura ( Diptera: Culicidae) and associated sylvan mosquitoes in south-
Science 110: 141- 142. of
site.
Entomol. 17: 453- 465.
Schulze, T. L.,
Howitt, B. F., H. R. Dodge, L. K.
Mansonia
R. H. Zimmerman, and J. D. Edman.
1980. Epizootiology of eastern equine encephaII. Population dynamics and vector potential of
Exterm. Assoc. 40: 141- 147. Hayes,
Morris, C. D.,
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L. E.
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outbreak.
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1- 10.
Am. J. Epidemiol. 87( 1):
Hagmann,
in New
description
and
The 1959
1968.
O. Sussman.
and
McNelly,
and
W. J. Crans.
1983. Epidemiological investigations of mos-
quito
populations
following
an outbreak of
eastern equine encephalitis among horses in New
Acad. Sci. 70: 320-327.
Jersey. J. Fla. Anti-Mosq. Assoc. 54: 21- 23. Morris,
C.
D.
evaluation
and of
S.
the
hypothesis eastern
transmission
of
lomeyelitis
Culiseta
Med.
Hyg.
by
1978.
Srihongse.
27: 1246- 1250.
transovarial
of
equine
melanura.
An
encepha-
Am. J.
Trop.
Veazey, J., D. Adam, and W. Gusciora. 1980. Arbovirus surveillance for eastern encephalitis in New
Jersey mosquitoes during the years 1960- 1969. Proc. N. J.
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Contr. Assoc. 67: 160- 171.
BULL. SOC. VECTOR ECOL., 11( 1):
SEPTEMBER, 1986
185- 190.
EVIDENCE INCRIMINATING COQUILLE7TIDIA PERTURBANS
DIPTERA: CULICIDAE) AS AN EPIZOOTIC VECTOR OF EASTERN EQUINE ENCEPHALITTSI
II. Ecological Investigations Following an Inland Epizootic in New Jersey W. J. Crans2, L. J. McCuiston2, and T. L. Schulze3
ABSTRACT: Ecological investigations following an epizootic of eastern equine encephalitis ( FEF) virus at an
inland location in New Jersey suggested that plant succession in a series of local man-made lakes had created adjacent breeding habitat for Coquillettidia perturbans and Culiseta melanura in areas where large numbers of horses were being raised. Data suggested that the ecological setting permitted Cq. perturbans to function as the primary vector to horses in areas where Cs. melanura was maintaining the virus at enzootic levels. Data also show that population levels of these two vector species change markedly from year to year and, thereby, regulate the risk of FF.F virus transmission in areas where the disease appears sporadically. INTRODUCTION
description of the Jackson Township area in relation
In 1983, four horse deaths due to
encephalitis ( EEE) virus were reported
from Jackson
region
epidemiological
where
Culiseta
horses
melanura
in that area
and
investi-
functioned
had
perturbans
Cq. Township
overwinter as adults,
Cs.
repeated vector
long
The
melanura
and
area.
the
study
following were
served as
locate the
Cs.
Since
both
extended well
Cq.
spring,
monitored
for
are presented
in this
have been
The combination of
abandoned.
region presents a unique ecological setting with small
rural farms interspersed with abandoned bog systems among typical pine barrens vegetation.
the
Location of Mosquito Breeding Habitat
in the
During September and October, 1983, the Jackson
species
Township focus was visited three times each week. All freshwater areas with emergent vegetation were
into the
perturbans
evidence of
the
of
year-
(
sampled for Cq. perturbans larvae using the modified bilge pump method described by Walker and Crans 1986). Aquatic plants that served as hosts for Cq. perturbans larvae were uprooted and brought back to
paper.
laboratory for identification. Lakes and bog systems were initially located by examining township maps. Later in the study, helicopter surveillance the
MATERIALS AND METHODS Description
Crans
systems
horse farming and cranberry farming in a pine barrens
breeding
melanura
involvement. The findings
investigation
industry is declining in that area and most of the bog
the
ecological
abated,
habitat
and
at
as the enzootic vector
investigations
winter months.
gations
Coquilletidia perturbans
were conducted to
Jackson
lakes. Most of the lakes were originally constructed to hold water for cranberry farming, but the cranberry
were affected and concluded that
primary vector to horses. Once the outbreak
of
contains an inordinate number of small, man-made
20
normally be Schulze ( 1986)
FFF.. Crans and
considered at risk to
km2 (
52
because they were limited to a inland focus that would not
conducted
breeding
small
a
unique
farms
common on small farms, backyard plots, and larger The area also and training facilities.
community in the pine barrens of Ocean County, New Jersey. The cases were
Township,
mil)
to the outbreak of EEE in horses during 1983. The region is noted for raising horses and horses are
eastern equine
of
and
located
the Area
Schulze ( 1986)
gave
a
complete
additional
wetlands
for
survey.
From
November, 1983, to February, 1984, the survey was
1New Jersey Agricultural Experiment Station Publication No. D-40101- 02- 86 supported by U. S. Hatch Act with partial support from the New Jersey State Mosquito Control Commission.
2Mosquito Research and Control, Cook College, Rutgers University, P. O. Box 231, New Brunswick, New Jersey 08903 USA. 3New Jersey
State Department
of
Health, CN- 360, Room 705, Trenton, New
Jersey 08625,
USA.
186
BULL. SOC. VECTOR. ECOL.
cuau.
Yw..«
w.+ x
S_
u" T'
a ••-&...
a,,
SEPTEMBER, 1986
y., t
S
x w"
s
t
w. .° awe°'
Y
.
.
Figure 1. Shallow man-made lake in Jackson Township, New Jersey showing emergent vegetation within the lake and Atlantic White Cedar( Chamaecyparis thyoides) in the background.
y
y
r
wig
t.
3i,
t I ,
j
4 .
rah
Figure 2. Dense
stand of
perturbans
Swamp
in Jackson
Loosestrife( Decodon
Township,
New
verticillatus),
Jersey.
the major host plant for Coquillettidia
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
187 s
3- ' 4" .,),..,, 4°,'
011?„ 4 ,
.
'
4
''. ..! 14-,$•....it '°
4..„.
.
4
b
1
1°
ate
1..tii•
4
',.'
IN
B'f
Via
Tk
m
s.
Figure 3. Aerial view of a typical man-made lake in Jackson Township, New Jersey showing proximity of: A) larval habitat for Coquillettidia perturbans, B) larval habitat for Culiseta melanura, and C) horse
pasture. (
x»
Photo courtesy of the Ocean County Mosquito Control Commission). r
d.,<
v
t #
4
b
fit
`
r
4. } ;
f'
1
b.
lit,'
J
4r, ?
r
1
r
of
s
\
i.
r
3 I
r
Figure 4. Sampling Swamp Loosestrife for Coquillettidia perturbans larvae with a modified bilge pump in late fall.
continued once
When ice
weekly.
covered
lakes,
the
designed using a heavy iron bit Areas were for ice fishing.
collections were made
to
SEPTEMBER, 1986
BULL. SOC. VECTOR. ECOL.
188
holes
out
chip
larvae and the results
perturbans
survey map Resting boxes were
trations of
Cs.
an
area,
breeding
a
the flooding of cranberry bogs. In those cases where aquatic vegetation had become established in the
the month of
littoral zone, Cq. perturbans larvae were found on the
adults were
melanura
Since Cs.
concen-
locate
to
during
adults
to
made
effort was
concerted
source.
constructed to function as reservoir water sources for
utilized
melanura
September. When Cs.
All but 2 of the 12 lakes examined were originally
were color coded on
the area.
of
a geodetic
found in
locate the
larvae
melanura
are pri-
pockets of water( Benzon marily found in underground 1984, Joseph 1984, McNelly 1984) the modified
bilge pump
Likely
as
used
again
was
areas of swampland
sampling tool.
a
containing
access
holes
for larvae
examined
underground water were
perturbans was primarily associated with the lakes.
Cq.
for
negative
or
positive
either
as
classified
remainder had been abandoned and served as low-lying Larval surveillance revealed that Cq. swampland.
to
grasses.
the
FEE
dry
ice
Cs.
and
resting boxes identified
Jersey
1
CDC All
once each week.
traps
15,
State Department described
by
of
Cq. with
from 25
specimens were
on a chill table and submitted to the
the methods
were
baited
sampled
were
melanura
New
Health for
Crans
virus assay by Schulze( 1986).
and
Twelve
under
lakes
man-made
located
during
cultivation
at
30 cranberry bogs
and
survey. Ten
the
time
the
TABLE 1 lists plants that were found to support
numbers of larvae. The Narrow-leaved Cattail Typha
of
of
the bogs were
the
study,
the
was a major host plant in the upper
angustifolia
reaches of the larger lakes. Cattail has been cited as a
typical host plant for Cq. perturbans in New Jersey swamps if the bottom substrate is loose enough to Swamp allow larvae to attach ( Hagmann 1952). Decodon
Loosestrife,
verticillatus
was
in the Jackson
area.
The plant was very common
along the margins of the shallower bog reservoirs and all of the stands that were sampled supported large numbers of larvae at the time of the survey. In some
a grass belonging to the genus Juncus
supported large numbers of larvae in very shallow areas close to the upland. Larval attachment to this plant was most common during the winter months when much of the lake surface was frozen solid.
Aquatic flora associated with Cq. perturbans larvae in the vicinity of
TABLE 1.
equine deaths due to Eastern Equine Encephalitis.
Shrub
Scientific Name
Common Name
Plant Type
Buttonbush
Cephalanthus accidentalis
Sweet Pepperbush
Clethra alnifolia Chamaedaphne calyculata
Leather Leaf
Swamp Swamp
Azalae
Rhododendron visosum
Loosestrife*
Decodon verticillatus
Arrowhead
Sagittaria latifolia
Narrow-leaved Cattail*
Typha angustifolia
Monocot
Rush*
Juncus spp.
Denotes
host- plant
Herbaceous
a major
found to
support the greatest numbers of Cq. perturbans larvae
lakes,
RESULTS
were
and the greatest numbers of Cq. perturbans larvae.
Cq. perturbans larvae during the survey period. Three
area to measure
October
to
were collected with
perturbans
Most of the bog systems supported some
contained the greatest amount of emergent vegetation
presence or absence of
test for the
June
and
primary reservoirs for abandoned bog systems
of the vector species one year after
From
virus.
plants,
groups of plants appeared to hold the greatest
In early June of 1984, adult collected from the Jackson Township
levels
herbaceous
associated with the lakes and lakes that served as the
and the
mosquitoes
outbreak and
shrubs,
rarely found in the bogs themselves. The larvae were
Adult Mosquito Collections
population
aquatic
emergent vegetation, but Cq. perturbans larvae were
to the survey maps of the area.
results were added
of
roots
at the time of the
survey.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
Larval Surveillance for Cs. melanura Larval surveillance in the Jackson
Cs.
that
melanura
light trap habitat the
that
The
thyoides.
lakes
contained
Cs.
provided
in
per 1, 000 specimens tested, the highest season rating for any of the collection sites that were being
south of the affected area
was
its tributaries.
and
Cq. Township
producing
cedar,
suitable
monitored in New Jersey that year( Crans unpublished data).
man-
The MFIR from August 24 to the end of the
perturbans.
season ( the period when virus was known to be
sustained a
present) was 7. 73. Although the figure is not as high
Chamaecyparis
as the 12. 93 MFIR obtained from Cs. melanura in 1983 ( Crans and Schulze 1986), data suggest that
zone of
subterranean
with suitable
Jackson the
with
behind the littoral
melanura
in
habitat
associated
white
cedar
seven
the end of the season. The minimum field infection
were
Atlantic
of
and
rate ( MFIR) for the entire season was 4.88 isolations
lake in Jackson
Nearly every stand
collection
than the
melanura
however,
lakes
made
Cs.
the
of
Township,
August 24
breeding
Toms River
of the
the
positive pools were recovered from that date through
widespread
located to the
was
drainage
Most
more
detected in area revealed
had indicated. Sizeable
were
collections
189
most
pockets
breeding
that
EEE virus was again at near epizootic levels during
habitat.
the 1984 season.
During the course of the survey, Cs. melanura were found
14
at
separate
locations in the Jackson
area.
In
CONCLUSIONS
most instances, the habitat consisted of a narrow band of cedar
swamp
of a man- made
that surrounded the
lake. While
very back portions in one area,
Data from the 1983 surveillance of equine deaths
not extensive
in Jackson Township, together with the follow-up
the numerous foci provided
local breeding habitat Township. In all but two and Cq. perturbans melanura larvae the same lake- bog system.
survey
throughout most of the
instances, Cs. were
found in
Adult
light
Jackson
Populations
traps.
CDC
1983,
averaged
During
trap
EEE
and
Virus
When the
traps
greater
series
of
results suggested that the species was abnormally
1984.
During
epizootic was
in
100
the
low
Cq.
the state over the same two year period confirm that
in 1983, Cq. perturbans
the entire
by
the
dry
occurring in locations
same
specimens
during 1984. Light trap records from other parts of
in
per
populations were at their
highest levels since 1975 ( New Jersey Agricultural Experiment Station year,
EEE
unpublished
claimed
35
horses
data).
Finley
During that 1976)
and
emergency surveillance efforts were instituted to
the peak population period of 1984, the traps
monitor the virus on a regular basis ( Crans et al.
only
The 84
eight specimens per night.
in 1984
were
melanura
collected
virus
was
detected
from the Jackson
in
site.
obtained.
Surveillance for the habitats utilized by Cq. perturbans
the
Cs.
TABLE 2
numbers of specimens that were tested and
F.F.F. isolations
1976).
tested for EEE virus
separate pools and none tested positive.
However, FEE
the
extremely
placed
then
that
were
specimens were collected
specimens collected
lists the
a
night.
averaged
in 13
indicated
collections
Township during
ice baited
that
high during the 1983 season and abnormally low
populations
only 84
season,
suggest
population data are available to determine comparative
in 1984 CDC
1984,
numbers of Cq. perturbans in that area, trapping
Mosquito
perturbans
in
epidemiological events may have accounted for the equine cases of EEE seen in that area. Although no
F.F.F. virus was first
and
epidemiological
Cs.
melanura
setting.
revealed
the two
vector species
involved.
New Jersey during 1984. No.
Period Entire Season
Enzootic Period* Period from date
of
unique
of man-made lakes that were originally created for cranberry culture had resulted in breeding habitat for Culiseta melanura
TABLE 2. Virus Isolations from Cs. melanura collected from Jackson Township, Collection
a
Plant succession in a series
No.
F.RE
Tested
Pools
Positive
1435
79
7
4. 88
906
46
7
7.73
first isolation ( August 24)
to end of season.
MFIR
190
SEPTEMBER, 1986
BULL. SOC. VECTOR. ECOL.
were present
along the
lakes,
and
band
narrow
formed
of cedar that
that had been dammed to create the were present on the vegeta-
Cq. perturbans
had
tion that
in the
streams
grown
result was adjacent
up along the lake margins. The habitat for the enzootic and epi-
Crans, W. J., J. D. Downing, and A. A. DiEdwardo. 1976. The emergency surveillance program in New Jersey for potential mosquito vectors of encephalitis
viral
Proc. N. J. Mosq.
in 1975.
Contr. Assoc. 63: 82-93.
zootic vectors of FFF.in areas where large numbers of horses
were
being
of
habitat
Township
in very
Since this type
raised.
throughout Jackson
was widespread
localized foci, virtually every horse farm in the area was in contact with the two major vector species. The MFIR for EEE in
1984
suggest
virus
the
that
high
represents a
probably
detected in 1983
and again
epidemiological
setting both
risk area whenever
Cram, W. J.
and
T. L. Schulze.
Coquillettida
incriminating
1986.
perturbans
Evidence Diptera:
(
Culicidae) as an epizootic vector of Eastern Equine Encephalitis.
I. Isolation of EEE virus
from Cq. perturbans during an epizootic among horses in New Jersey. Bull. Soc. Vector Ecol. 11( 1):
178- 184.
vector species are present in high numbers. Since no human of risk
in
cases were reported
to
humans
cannot
either year,
be accurately
the
question
Hagmann, L. E. studies
assessed.
Mansonia perturbans recent
1952.
in New
Jersey.
Proc. N. J. Mosq.
Exterm. Assoc. 39: 60- 66. Acknowledgements
The Control
Ocean
thank the
authors
Commission for portions
ecological
their
of this
County Mosquito during the
support
Mr. Tom
investigation.
Candeletti and his staff were the first to identify Swamp Loosestrife as a major host plant for Cq. Candeletti
Romanowski
Michael
perturbans.
spent countless
and
hours in the field
Richard with
the
Finley, J. E. Jersey in
Arbovirus activity in New Proc. N. J. Mosq. Contr.
1976.
1975.
Assoc. 63: 9- 13. Joseph,
S.
1984.
R.
copter
time to
survey
maps.
locate
wetlands
also
that
provided
did
Appreciation is also
heli-
not appear on
extended
to Dr.
melanura
in
25- 32.
authors selecting sampling sites. The Ocean County Mosquito Control Commission
Culiseta
Maryland. Proc. N. J. Mosq. Contr. Assoc. 71:
McNelly, J. southern
Culiseta
in
R.
1984.
New
Jersey. Proc. N. J. Mosq. Contr.
melanura
Assoc. 71: 33.
James Westman, Professor Emeritus, Rutgers Univerin sity. Dr. Westman, an aquatic biologist, assisted the initial survey of Jackson Township and conceived the idea of using a bilge pump to sample a mosquito species
that attaches to
the root
system
of aquatic
W. J. Cram, and W. E. Parkin.
1984. Multi-agency response to Eastern Equine Encephalitis
in
Jackson
Township,
Ocean
County, New Jersey. Proc. N. J. Mosq. Contr. Assoc. 71: 87-93.
plants.
REFERENCES CTlED Benzon, G. L. melanura
Proc.
Schulze, T. L.,
N. J.
1984. (
Larval habitat
Walker, E. D. of
Culiseta
Coquillett) in Eastern Massachusetts.
Mosq.
Contr. Assoc. 71: 23- 24.
and
W. J. Cram.
1986.
A simple
method for sampling Coquillettidia perturbans
larvae. J. Am. Mosq. Contr. Assoc. 2(2): 239240.
BULL SOC. VECTOR ECOL., 11( 1):
191- 193.
SEPTEMBER, 1986
IMPACT OF TWO UNICELLULAR ALGAE ON THE DEVELOPMENT OF CULEX QUINQUEFASCIATUS SAY( DIPTERA: CULICIDAE) Lal S. Mianl and Mir S. Mullal
ABSTRACT:
bijugatus)
Studies
out on the
carried
effect
of two
algae (
Kirchneriella irregularis
and Scenedesmus
on the development of 1st-instar larvae of Culex quinquefasciatus Say through adult emergence
showed that the development of mosquito larvae exposed to 1, 5, and 25 percent culture of both microphytes in the presence of larval food was not significantly affected. None of the larvae in both control and algal treatments
in the absence of larval food developed into pupae. The survival of larvae, however, was directly related to algal concentrations, thus, suggesting that both algae were utilized as an inadequate source of food by the starving larvae.
27± 1° C).
INTRODUCTION A
substantial
amount
of evidence
mosquito
Examples
species.
the
supports
observations that various algae are toxic to of algae
different that
species
have demonstrated larvicidal activity against several species of Aedes, Anopheles, Culex spp., etc.
include Chara
spp. ( Caballero 1919, Frederici 1927, Castillo 1944), Cladophora spp. ( Hamlyn-Harris 1928, Amonkar 1969), Spirogyra sp. ( Boyd and Foote 1928,
1956), Schizothris sp., Hapalosiphon
Chlorella
1974),
hieroglyphicum 1981).
1932), Anabaena spp. ( Gerhardt sp. ( Amonkar 1969), Microcystis
Bradley
sp. ( Ilyaletdinova ellipsoidea
and
( Dhillon 1980, Dhillon Marten ( 1984)
Recently,
Kirchneriella - like
phytoplankton
Dubitskii
Rhizoclonium
and
and
Mulla that
reported effective
was
in
In each treatment 20 1- day-old 1st- instar
larvae of the test mosquito were added to 200 ml of diluted algal culture of deionized water in a 300 ml Pyrex® glass bowl. In two concurrent tests, one set received 10 mg of larval food( chicken mash:Brewers
yeast, 3: 1) daily and the other set was maintained food.
Controls in these tests included mosquito larvae in deionized water with and without larval food. All treatments and controls were repliwithout the
cated four times. Daily observations on larval and pupal
mortality
were
continued
through
adult
emergence or until termination of the test, maximum
70 days. The above sets of experiments were also repeated later at a different time. Data generated on
these tests were statistically analyzed and comparison
of means was done accordingly, using Duncan's multiple range test( DMRT).
suppressing larval populations of Aedes albopictus. Pure and
and
diluted( 50%)
Anopheles
Marten 1984,
unpublished
data
latter two
algae on
Culex
of
Anopheles
albimanus
in the
laboratory
quadrimaculatus
laboratory
on the
data). This
impact
of
dilute
the growth and
quinquefasciatus
Say
RESULTS AND DISCUSSION
were also found to be
lethal to the 1st- instar larvae and
both K. irregularis
cultures of
Scenedesmus bijugatus
The results of tests on the survival of Cx. in different algal cultures are
quinquefasciatus
paper persents
presented
cultures of the
survivorship of this mosquito in cultures of K. irregularis showed that the mean survival based on
development
of
larvae.
net
adult
in TABLES
emergence
1
was
and
2.
Studies on the
higher ( 91%)
in controls
than in different algal treatments ( 76- 88°, TABLE 1). MATERIALS AND METHODS Pure cultures of Kirchneriella irregularis and Scenedesmus bijugatus tested were at concen1, 5, and 25 percent of pure culture in rations, deionized Culex
water
for their impact
quinquefasciatus
through
adult
iDepartment
eclosion
of
Say under
on the
survival of
from the larval
laboratory
Entomology, University
of
stage
conditions
The variability between treatments of test I and II seems to be due to day-to-day variations since both tests were carried out at different times with mosquito
larvae from different batches. In the absence of larval food, larval development was delayed in either test; the larvae in test I were unable to pupate even in 70 days. Where, in the no- food control, larval food was
the limiting factor affecting larval growth and develop-
California, Riverside, CA 92521- 0314 USA.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
192
TABLE 1. Survival of Culex quinquefasciatus in the presence of Kirchneriella irregularis.a Percent survival with and without food at posttreatment intervals( days) in Test I and II Algal Culture%
5 25 Control
83ab 80a
76a
91b
II
I
II
I
88a
d
96a
16a
85a
I
II
76a
25
35
55
70
I
I
I
I
II
-
61a
13a
5b
3a
0
38a
13a
8a
3
95a
54b
81a
-
78a
-
91a
49b
69a
-
56a
48a
15a
4a
1
-
Ob
14a
Ob
Ob
1l
9b
3a
3
-
88a
19
17
12
17 I
1
No food°
foodb
Larval
91a
-
aMean of four replications with 20 1st-instar larvae per replication. Means followed by same letter(s) in
a column are not
significantly different from
one another(
DMRT, p= 0.05).
bFigures show percent adult emergence. All figures represent larvae which did not pupate.
dData were not recorded.
TABLE 2. Survival of Culex quinquefasciatus in the presence of the alga Scenedesmus bijugatus.a Percent survival with and without
food at posttreatment interval( days)
Algae
No Food°
Foodb
Culture(%)
19
17
17 TEST I
TEST II
TEST ll
1
83a
80a
11a
8
5
91
a
84
a
25 b
16
25
96
a
88
a
35 b
33
92
a
85
a
0
Control
a
0
aMeans of 4 replications with 20 larvae per treatment.
Means followed by same letter in a column are not significantly different from each other( DMRT, P : 105).
bFigures in these columns) show adult emergence. cAll figures
represent
larvae
which
did
not reach pupation.
SEPTEMBER, 1986
the
ment,
BULL. SOC. VECTOR ECOL.
survival
cantly higher
of mosquito larvae was signifilasted longer in algal treatments This indicates that the algae act as a
and
than in controls.
food for the
source of
food
this
source
may
larvae. However,
developing
not
be
sufficient to afford corn-
193
Boyd, M. F.
and
bionomics
H. Foote.
of
1928.
American
Studies on the The
anophelines.
alimentation of anopheline larvae and its relation
to their distribution in nature. J. Prevent. Med. 2: 219-242.
plete larval development through adult emergence. Tests on the survival of Cx. quinquefasciatus larvae on cultures of S. bijugatus yielded data almost similar to those obtained with K.
irregularis.
Bradley, G. H. 1932. Some factors associated with the breeding of Anopheles mosquitoes. J. Agric. Res. 4: 381- 399.
In both test I and II when larval food was provided, the mean survival of mosquito larvae through adult eclosion
was
not
significantly
treatments as compared
by
reduced
algal
the control( TABLE 2).
with
hi the absence of larval food, the surviving larvae being
however, lived longer in the
unable to pupate,
in the
algal cultures than
This
control.
also suggests
Caballero, A.
1919. La Chara foetida A. Br. y las
larvas de Stegomyia, Culex
y Anopheles.
Bol.
Sol. Esp. Hist. Nat. 19: 449- 455. Castilla, R. L.
1944.
The possible role of Chara
fragilis in mosquito control. Science 100: 266.
that the algae were the only source of nourishment inadequate for
yet
larvae
the
to
complete
their
development. Under ample
present
food
of
irregularis
nature, there
is
an
and suspended organic solids
in larval habitats.
The
both K.
presence of
and S. bijugatus at 1, 5, and 25 percent
dilutions in habitat larval
in
eutrophic conditions
quantity
mosquito
water
not
may
significantly
In
populations.
an
reduce
earlier
study,
Marten ( 1984) reported that an abundant growth of Kirchneriella spp. ( or Chlorella) in outdoor gallon
water jars was responsible for the suppression of larval development
of
Ae. albopictus.
Later, Marten
1984, unpublished data) found that pure
50%)
cultures
bijugatus
of
were
An.
albimanus and
both
K.
lethal to
and
diluted
irregularis
and
lst-instar larvae
of
In these experiments, however, the very small 5 larvae/ cup) and lack of replications
tory.
sample size (
make
his
presence algae
in
such
actual
high test concentrations
field
is
situations
of
both to
subject
speculation.
Entomology,
of
Kirchneriella irregularis
Scenedesmus bijugatus
were
G. Marten,
Hawaii.
University
of
kindly
provided
and
by
Dr.
Dhillon, M. S.
1969.
Freshwater
of
Calif.,
and
M. S. Mulla.
1981.
Biological
activity of the green alga Chlorella ellipsoidea
against immature stages of mosquitoes. J. Am. Mosq. Contr. Assoc. 41: 368- 372. Frederici, E.
1927. L'azione tossica delle Charae sulle larva dei Culicid' e. Redia 16: 17- 28.
Gerhardt,
R.
W.
1956.
Present knowledge concerning the relationship of blue- green algae
and mosquitoes in California rice fields. Proc.
Calif. Mosq. Contr. Assoc. 24: 47- 50.
Hamlyn-Harris, R. 1928. The relationship of certain species of algae to breeding places of mosquitoes
llyaletdinova, S. G.
Bull. Entomol. Res. 18: 377-
and
A. M. Dubitskii.
1974.
Toxicity of Hapalosiphon fontinalis, Anabaena variabilis
and
Microcystis
aeroginosa
with
respect to larvae of blood-sucking culicide. IZV.
REFERENCES CITED Amonkar, S. V.
Univ.
Riverside, Calif., 120 pp.
389.
Acknowledgements The
Dept.
in Queensland.
pure cultures of
Biological activity of natural
Moreover, the
observations questionable. of
1980.
products of algae and a hydrophyte against mosquitoes. Unpubl. Ph.D. Disser- tation,
S. An.
in the labora-
quadrimaculatus
Dhillon, M. S.
Akad. Nank. Kaz. SSR. Ser. Biol. algae and their
12: 29- 35.
In Russian).
metabolites as a means of biological control of mosquitoes.
Unpubl.
Ph.D. Dissert.,
Entomology,
Univ.
Calif., Riverside, Calif.,
102 pp.
of
Dept.
of
Marten, G.
1984. The impact of planktonic flora and fauna upon Aedes albopictus larvae. Bull.
Soc. Vector Ecol. 9: 1- 5.
SEPTEMBER, 1986
194- 196
BULL. SOC. VECTOR ECOL., 11( 1):
SURVIVAL AND OVIPOSITIONAL RESPONSE OF
CULEX QUINQUEFASCIATUS SAY( DIPTERA: CULICIDAE) TO SEWAGE EFFLUENT Lal S. Mianl and Mir S. Mullal
Studies on the development and survival of immature stages of Culex quinquefasciatus Say in
ABSTRACT:
secondary sewage effluent ( SSE) under laboratory conditions showed that the development of larvaeLarval was larval food. in SSE significantly lower in SSE than
under
or
control
conditions
provided
with
development in SSE with or without larval food was also delayed, indicating that besides certain nutritional
factors lacking in SSE, there was some other factor(s), the lack or presence of which was detrimental to larval
In addition, SSE was highly attractive to ovipositing female mosquitoes under laboratory
development. conditions.
INTRODUCTION
depending
on the age of the
conducted
in 300
ml
SSE
of
or
ml
larvae.
The tests were
Pyrex® glass bowls with 200
deionized
water.
Each bowl was
sewage
stocked with 20 1- day-old 1st-instar larvae of Cx.
SSE) at the Riverside Wastewater Treatment Plant were observed to support the development of large populations of Chironomus decorus Johansen,
quinquefasciatus. Each treatment was replicated four
floating
Observations on the development of larvae in each
These basins were 10 egg- rafts was never recorded. feet deep and the SSE level ranged between 5 and 8 on the daily sewage inflow into feet
treatment and control were made daily and larval food where necessary was added. The data thus obtained
basins
Equalization
holding secondary
effluent(
but
larvae,
of mosquito
evidence
or
pupae,
times. The bowls were kept in a room where the
The basins
system.
SSE from these habitats the
mosquito,
tory
Culex
the
of
apparent
for its
was tested
quinquefasciatus
maintained
at
26±
1° C.
was evaluated using the Duncan's multiple range test
floating (
DMRT).
the
In the oviposition test, two 112 ml wax paper
effect on
cups, one containing 100 ml of deionized water and
in the basins,
ovipositional
and
survival
have any
not
breeding
of mosquito
absence
did
Because
vegetation.
or
material
also
was
were analyzed statistically and comparison of means
deep, depending
the
temperature
ambient
the
response
of
Say
labora-
under
the other having 100 ml of a 2- day-old SSE with bottom x
conditions.
22. 9
mud( x
10%),
22. 9
were placed in a screen cage( 30.5
cm).
The cups alternated in position
in the cage. Then, 20 1- week- old gravid females of
Cx. quinquefasciatus were released into the cage. The
MATERIALS AND METHODS
test was replicated six times at a room temperature of Studies mosquito
to
out
carried
were
larvae in SSE
ovipositing
of
survival
the attractancy of SSE
and on
mosquitoes
the
on
in the
after 13 hours of scotophase.
To
laboratory. out
carried
24-27° C. Observations on oviposition were recorded
RESULTS AND DISCUSSION
at
study survivorship, two tests were The first test, conducted from different times. October through November, 1984, consisted of
The results of survivorship studies showed that
SSE
mean survival of larvae through adult emergence in
The
SSE
exposure with
and
second
of mosquito without
larval food,
test ( Nov.
provided containers
10
food, SSE
with
basin,
control
and
larvae in
1984 of
SSE
percent with
containers
and
Jan.
of
controls.
1985),
however,
and
mud
without
to
1Department
larvae
of
at
10- 15
mg
Entomology, University
of
was not significantly 93%, TABLE 1).
control (
Nevertheless, mosquito survival in SSE alone ( 23%
larvae) was significantly lower from survivorship data in control. The 4th-instar larvae surviving in SSE
larval food.
per
of
larval
Larval food ( Lab. Chow: Brewers Yeast, 3: 1) provided
larval food ( 84%)
from the
with and without
bottom
with
different from that
was
container,
without food were weak, sluggish, or moribund at the end of the test period.
These data indicate that the
California, Riverside, CA 92521- 0314 USA.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
195
TABLE 1. Survival of Culex quinquefasciatus in secondary sewage effluent( SSE). a Mean%
SSE
survival by stage in
SSE
with
food
Control with food
Interval
Total
Ab
Days)
L
0
100
0
0
100
0
0
100
0
0
6
95
0
0
98
0
0
100
0
0
8
90
0
0
98
0
0
90
10
0
10
90
0
0
20
78
0
10
80
10
P
L
P
A
L
P
A
14
85
0
0
0
20
78
0
10
90
20
60
0
0
0
0
84
0
0
93
24
38
0
0
0
0
84
0
0
93
29
23
0
0
0
0
84
0
0
93
23a
0
0
0
0
84b
0
0
93b
Yieldc
Mean of 4 replications with 20 larvae per treatment.
bL=
larvae, P=
pupae, and
A=
adults.
Means followed by the same letter in the last row are not significantly different from one another(
DMRT, P= 0.05).
TABLE 2. Survival of Culex quinquefasciatus in secondary sewage effluent( SSE). a Mean% SSE
SSE
Interval
w/ bottom
mud(
Days)
L
P
A
L
SSE
10%)
P
survival by stage in w/food
Control
A
L
P
Dist. Water
w/food)
no mud)
A
L
no food)
P
A
L
P
A
0
100
0
0
100
0
0
100
0
0
100
0
0
100
0
0
10
8
0
0
65
0
0
59
0
0
69
24
0
23
0
0
20
14
0
0
45
4
0
0
0
0
0
0
93
15
0
0
30
0 0
0 0
22
0
1
0
0
0
0
0
93
9
0
0
38
6 6
12
1
6
0
0
0
0
0
93
9
0
0
42
3
0
0
12
1
6
0
0
0
0
0
93
9
0
0
Total Yield( L,P, A)°
3a
19c
56d
93e
aMean of 4 replicates with 20 larvae per replicate.
bL= larvae, P= pupae, and A= adults. Means followed by different letters in the total yield row are significantly different from one another(
DMRT, P=
0. 05).
9b
196
suppressant effect of
be due to the
could
be
SSE
food, this
delayed
by
explained
The
data
food,
lower adult
could
be
date.
day- to- day
biochemical
the physicochemical or
SSE
SSE in both tests
since the
The data in TABLE 2
information of which
on
did
mud, the net
mud, the presence
larvae that
with
bottom
12% larvae, 1%
significantly lower
marginal
vegetation
in
Cucamonga
SSE aquaculture ponds with a dense cover of water
Comm.) and in San Diego ( B. A. Wilson 1983, Pers. Comm.).
Data from the oviposition test showed that SSE was
highly
mosquitoes.
to female attractive ovipositing The mean ovipositional response to
SSE was 17. 3 egg rafts/ cup as compared with zero egg
rafts
in
control.
Whether this substance or its
odor emanating from
attractive to laboratory environment can work with the same efficiency out in ovipositing
mosquitoes
SSE being in
enclosed
the field, is subject to further investigations.
In our observations on the absence of egg rafts
with
in three equalization basins, it might be possible that
pupation and adult emergence as
the physical features of the basins were not conducive
data in SSE
adults) and control(
from
percent (
adults) was
in
the same
also yielded additional
bottom
10
of
constituents
However, in SSE
adults.
delay
compared with
that apart
with
yield of
6%
considerable
SSE
larval
fluctuations in
was not of
with
Sjogren and Mulla 1968, Mulla et al. 1970) and in
larvae in
with
provide nourishment to some
developed into
pupae, and
to those of test 1
in SSE
(
hyacinth in Roseville ( K. R. Townzen, 1983, Pers.
survival of
emergence
attributed to
and adult
control.
conducted a month
almost similar
However, the lower
TABLE 1). and
test( TABLE 2)
second
yielded
phenomenon
pupation
data in
emergence as compared with
later
lagoon
lack
larval
with
several other species, was reported in raw sewage
larval development may of certain factor( s).
on
presence or
Even in SSE
SSE
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
93%
with
adults).
larval food ( 56%
to oviposition by female mosquitoes.
These data suggest
certain nutritional elements
lacking
in
REFERENCES CITED
SSE, there could have been other environmental physicochemical
vegetation,
of mosquito
factor( s),
the lack of
breeding
such
in SSE
marginal
as
which resulted
in the
equalization
absence
basins.
Usually the presence of vegetation in raw sewage or even SSE can offer ideal breeding habitats for mosquitoes capable of breeding in polluted waters. In different parts of California, for example, the breeding of various species, quinquefasciatus,
such as
Culex
Culex tarsalis
peus
Speiser, Cx.
Coquillett,
and
Mulla, M. A., H. A. Darwazeh, and D. R. Peters. 1970.
Mosquito control in sewage oxidation
ponds with
drip
and pour- in
larvicides.
Mosq.
News 30: 457- 460. Sjogren, R. D. application
and of
M. S.
Mulla.
three
1968.
Drip
organophosphorus
insecticides for mosquito control. Mosq. News 28: 172- 177.
BULL. SOC. VECTOR ECOL., 11( 1):
197- 198.
SEPTEMBER, 1986
FIELD TESTS IN WESTERN MONTANA OF
MECHANICAL TRANSMISSION OF TULAREMIA BY BITING FLIES DIPTERA) BETWEEN IMMOBILIZED LABORATORY ANIMALS C. B. Philipl and W. L. Jellison2
Tularemia is nature
bacteriologic
to
disease in
enzootic "
man"
by
caused
Francisella tularensis
agent,
hemisphere
northern
This
interesting,
an
communicable
As potential vectors, species of the above three families of biting Diptera were tested in the field as
the
in
the
seasonal and flying conditions made the flies
the New and Old Worlds.
of
available.
memorializes one of the
early investigators of biting fly involvement, Dr. Edward S. Francis. G. W. McCoy, with C. W. Chapin, named the agent of a " plague- like
disease
of
rodents"
he
that
healthy test guinea pigs were transported in protected cages to the test sites for exposure on immobilization
had
spread-boards. The primary objective was to check
discovered a year earlier in Tulare County, California. There is not the time, nor is this the place, to review the versatile methods
naturally
deer
reported was as" attributed
Francis
to
and
by but
propagates,
fly
bites
the
transfer
agent
by Jellison(
1974).
Other
Each fly in such test bitings was then also confined in a clean tube for storage at lowered
species
Tabanidae were reported by him to be naturally infected. It became obvious to investigators that interrupted)
by biting flies bitings
adaptation to the
fly
rather
biting
temperature of 16° C to reduce activity until further delayed test feedings were attempted on return to the
mechanical
biological
laboratory, the period determined by the survival vigor of the flies during storage. They were warmed
Montana,
further feedings on new healthy guinea pigs.
vectors.
In the Bitter Root certain
by by
was
than
to room temperature to attempt stimulation of delayed
Valley
species of
of western
Tabanidae ( horse and deer and Ceratopogonidae
Injection"
flies), Simuliidae ( blackflies),
in the field
midges) were tested
by
the authors to
simulate under more natural conditions rather than artificial,
multi-
laboratory ways the
among
haematophagus
cages.
disease
This is
agent
is
one
considered to
species that were
locally
While
strains of
laboratory
restrictive
flies were also collected in saline diluent in tuberculin syringes and applied with the needles removed to the abraded skins of fresh animals.
be
propagated
available
hosts
F. tularensis have been
of
low
and
was
shown
by
those
to
be
characteristic
tularemic
pathology
observed
at
necropsy as described by Davis et al. above and Philip and
virulence
reported
authors
Typical infections" in test animals refers to
of
for test.
in Montana Davis et al. 1934), the " B. C. Strain" used in the present tests was from an isolate in British Columbia animals
Davis et al. 1934) of nearly or freshly dead, stored
hypothesized
MATERIALS AND METHODS
for
(
single flies. Fresh fecal droplets for test from stored
Diptera of the
of flies refers to intraperitoneal
inoculum prepared by trituration in standard saline
in
rabbits and other susceptible native
biting fly
on
to be used should an uncontrollable fly swarm occur.
of
transmission
presumably
captured, and precautionary cover at hand did not have
by subsequently by
and
organisms,
that became available during a given test. Every fly
Utah first
northern
tularemia
visitor that attacked the donor animal fortunately was
Chrysops discalis
of
of
contaiminated mouth parts of the different fly species
the earliest ways
of
fever" in
Mayne ( 1921),
others as reviewed
disease
this
which
one
Locations of suitably abundant pop-
ulations were predetermined and donor( infected) and
of
virulence.
high
Williams ( 1985).
In tests deemed crucial,
infection was further confirmed by culture of blood from moribund animals, and by spleen vaccination of a second passage guinea pig ( Philip et al. 1932). Moribund donors were always necropsied after return to the laboratory to confirm their tularemic in-
fections. Tests which resulted " negatively" indicated no signs of disease in a given test animal in which
1Paper presented at the Pacific Division Meetings, American Association for Advancement of Science at the University of Montana, Missoula, 11 June, 1985.
2California Academy of Sciences, Department of Entomology, Golden Gate Park, San Francisco, California, 94118. 3Rocky
Mountain
Laboratory(
retired),
Hamilton, Montana, 59840.
SEPTEMBER, 1986
BULL. SOC. VECTOR ECOL.
198 case
the
blood
animal' s
serologically for
was checked
DISCUSSION
anti- tularensis agglutinins ( Philip et al. 1932) before As is well-known, maximum bacteremia occurs in tularemia infected animals during the terminal stages of the disease. Hence, donor guinea pigs taken
was considered" negative."
the test
RESULTS blood
interrupted
Undelayed
between
meals
guinea pigs resulted
in
into the field for interrupted fly bitings in these tests
transfer of
healthy F. tularensis by Hybomitra rupestris in 3 of 4 in the and by H. osburni attempts; by H. frontalis noctifer pertinax Chrysops only test made of each; by in 1 of 7; and by Simulium decorum katmai in 3 of infected
14
and
were
10
15
of
delay
while
1
2
24 hours in tests
of
of
minutes
The data here presented suggest that other
and
7
of
2 delays
C.
of
24 hours, 7
of
plus
48 hours
of
2
and
in the aggravation of eipzootics among susceptible
96
of
domestic and native animals, or in occasional human
negative
infection as well.
involved bitings of 2 flies;
pertinax
injected 10
13 days later into
and
separate
REFERENCES CITED
animals, each of which became infected. One of the concerned a fly negative 24- hour tests of S. d. katmai to harbor F. tularensis proved which was by One
injection 4 days later.
injected into a test
when
Davis, G. E., C. B.
Philip,
and
R. R. Parker.
1934.
The isolation from the Rocky Mountain wood tick (Dermacentor andersoni) of strains of Bact.
of the ceratopogonids was
for 9 days
to have retained the organisms
also shown
previously incrimi-
Chrysops discalis,
to the natural incidence of tularemia. The widespread occurrence of this disease in the northern hemisphere emphasizes the potential involvement of biting flies
ceratopo-
S. d. katmai. The 11- hour
of
n.
these were
H. frontalis,
to
nated by Francis and Mayne ( 1921), may contribute
and
hours in tests test
addition
followed positively. Negative results 11 hours in tests of C. n. pertinax
gonids, resulted
delays:
with
rupestris
undetermined
of
Their characteristic
tests.
30 hours
of
and
minutes
condition.
species of biting flies, particularly of tabanidae, in
H.
refeedings with stored
Two delayed
moribund
confirmed the validity of the respective transmission
others.
involved intervals
in
lesions, checked at later necropsy in the laboratory,
tularense
of low virulence for guinea pigs and
domestic rabbits. Am. J. Hyg. 19: 449- 456.
animal.
Injection of suspensions of single H. rupestris into
1 to 4 days
guinea pigs, each
blood
also
meals
instances.
Similar tests
days,
resulted
also
of
after
their
initial
in 4
infection
produced
of
5
the H. frontalis, stored 11
9- day H.
of
positively;
osburni
resulted positively in 1 of 2 attempts; and of 3- to 44day
C.
stored
Results
attempts.
were positive
n. pertinax
injections
of separate
in 16 to 18
20 S. d.
of
katmai after a few hours to 9 days storage were 5
positive
The
times and negative species of
to
amenable
deleterious
flies
cage of
effect
under experiment were not
and
ingested F.
tularensis on their
infected C.
proven
longevity
could not
ticks, unpublished)
However, 3
possible
the
conditions
to reduced
survival( comparable
some
15 times.
be
observed
in
ascertained.
lived
n. pertinax
as
long as 28, 33, and 44 days in chilled tubes. Feces lets in
of
3 C.
storage tubes
original
blood
suspensions
separate
2, 4, 25,
and
produced
meals
were
collected as
n. pertinax
rubbed
on
the
Eleven
test guinea pigs.
fresh
26 days
infection abraded
drop-
after
the
when
skins
other similar
of
fecal
tests at 2 to 18 days were negative. In 3 fresh fecal
tests
days, from H. frontalis stored 7 days, osburni and
the
other
from H.
stored
the
2 positively.
rupestris
stored
4
11 days, and from H.
first
resulted
negatively
Francis, E.
and
B. Mayne.
1921.
Experimental
transmission of tularemia by flies of the species Chrysops discalis.
Publ. Hlth. Rpts. 36: 1738-
1746. Jellison, W. L.
1974.
Tularemia in North America
1930- 1974. Univ. Montana, Missoula, 276 pp. McCoy,
G. W.
and
C. W. Chapin.
1912. Bacterium
tularense
the cause of a plague- like disease of U. S. Public Health Service, Publ. rodents. Hlth• Bull. 53: 17- 23. 1932.
Philip, C. B., G. E. Davis, and R. R. Parker. Experimental transmission of tularemia
by
mosquitoes. Publ. Hlth. Rpts. 47: 2077- 2088.
Philip, C. B. and W. L. Jellison. 1985. Tests in the field
in
western
Montana
of
mechanical
transmission of tularemia by biting flies between immobilized laboratory animals. Proc. Pac. Div. Am. Assoc. Adv. Sci. 4: 38.
Philip, C. B. and S. C. Williams. 1985. Tularemia and other problems in livestock induced by wood ticks in Montana. Bull. Soc. Vector Ecol. 10( 1): 45- 47.
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