Beruflich Dokumente
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Bulletin of the
SOCIETY FOR
VECTOR ECOLOGY
4" 4
JIL
December, 1993 f
BULLETIN OF THE
SOCIETY FOR VECTOR ECOLOGY
Volume 18- Number 2 December 1993
EDITORIAL BOARD
Mir S. Mulla, Chairman
University
of
California
W. J. Crans
R. S. Nasci
Rutgers
CDC-DVBID
University
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R. C. Axtell
C. I. Dahl
University
University
Fort Collins, CO
M. W. Service
of
Uppsala
Raleigh, NC
Uppsala, Sweden
Liverpool, England
N. Becker
F. R. Holbrook
E. D. Walker
USDA- ARS
Laramie, WY
D. R. Barnard
R. S. Lane
S. K. Wikel
University of California
Berkeley, CA
USDA, Gainesville, FL
H. Briegel
G. G. Marten
S. C. Williams
New Orleans, LA
San Francisco, CA
E. P. Catts
L. S. Mian
University
Germany
University
of
of
Heidelberg
Zurich
Washington State
Stillwater, OK
University
Pullman, WA
San Bernardino, CA
Sustaining
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BOARD OF DIRECTORS
1993 OFFICERS
President
Norbert Becker
President- Elect
Susan M. Palchick
Vice- President
Bruce F. Eldridge
Past- President
William K. Reisen
Secretary- Treasurer
Gilbert L. Challet
REGIONAL DIRECTORS
Southwestern
Minoo B. Madon
Northwestern
Steven V. Romney
North Central
Thomas R. Wilmot
South Central
Cluff E. Hopla
Northeastern
Wayne J. Crans
Southeastern
Jonathan F. Day
European
Frantisek Rettich
BULLETIN
OF THE
December, 1993
Volume 18
CONTENTS
ii
ix
1993)
1993)
xi
xii
Book Review
Arthropods of Medical and Veterinary Importance: A Checklist of Preferred Names and Allied Terms. A.
R.
Reviewed
Pittaway
by
D. L. Baumgartner
75
Proceedings
7th SOVE Annual European Branch Meeting, Bologna, ITALY, 25 August 1992
Tick Ecology and the Search for a Possible Lyme Disease Spirochaete in Southeastern Australia
Richard C. Russell, S. L. Doggett,
and
R. Munro
77
First International Congress of Vector Ecology, San Diego, California, 4 October 1993
The Role
of
Ecology
and
Ecological
Modeling
M. W. Service
in Vector Control
85
Submitted Papers
Borrelia burgdorferi Infection in Ixodes scapularis ( Acari: Ixodidae) in Kent County Maryland
F. P. Amerasinghe
and
T. W. Scott
99
Size of Emerging and Host Seeking Aedes aegypti and the Relationship to Containers and Blood-Feeding
D. D. Chadee
105
Daily Survivorship and Life Span of the Mosquito Eretmapodites quinquevittatus Theobald ( Diptera:
Culicidae) Under Laboratory Conditions Compared toAedes aegypti,Aedes abopictus, andAedes bahamensis
D. Vodopich
109
J. D. Lang
County, California
Elevated Dosages of Bacillus thuringiensis var. israelensis Fail to Extend Control of Culex Larvae
M. S. Mulla, J. D. Chaney, and J. Rodcharoen
114
A. M. Helleck, W. K.
Fleas Found
on
Blow
and
Invading
Hartberg,
Fly
of
Fly,
Chrysomya
megacephala (
A. R. Olsen, T. H. Sidebottom,
and
125
as an
S. G. Bennett
133
Preliminary Research on Bacillus thuringiensis var. israelensis Use for Mosquito Control in Northeast
K. Ouzounis
Greece
and
A. Samanidou- Voyadjoglou
147
and
M. S. Mulla
152
Field Trials of a Fizzy Tablet With Bacillus thuringiensis subsp. israelensis in Forest Spring Ponds in
A. G. Eriksen
160
Occurrence and Abundance of Cyanobacteria in a Brackish Marshland and Their Ingestibility by Mosquito
I. Thiery, G. Sinegre, and N. Tandeau De Marsac
Larvae
164
O. Skovmand
Denmark
and
Water Depth and Larval Density Affect Development and Accumulation of Reserves in Laboratory
Populations
of
Mosquitoes
S. E. Timmennann
and
H. Briegel
174
The Bulletin of the Societyfor Vector Ecology is an international journal concerned with
all aspects of the biology, ecology, and control of arthropod vectors and the interrelationships
between the vectors and the disease agents they transmit. Thejournal publishes original research
articles and research notes, as well as comprehensive reviews of vector biology based on
presentations at Society meetings. All papers will be reviewed by at least two referees who are
qualified
suitability for
publication.
Acceptance of
manuscripts is based on scientific merit and is the final decision of the editor, but these decisions
may be appealed to the Editorial Board.
encouraged.
Wordstar formats are acceptable, as well as unformatted text files. Please indicate the type of
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a separate page, in order: Title page, abstract, text, acknowledgments( ifappropriate), references
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and their affiliations and the
identification
corresponding
author.
Pages
should be numbered consecutively starting with the title page. References should conform to
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Page charges, which partially defray the cost ofpublication, are$ 35 per printed page.
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ii
NUMBER 2
DECEMBER, 1993
VOLUME 18
IN MEMORIAM
rh
4 nRnx
was
Thomas D. Mulhern,
devoted
control.
and
his lifetime
M,"
such was
who throughout
dedicated to
Tommy
the case
with"
mosquito control
he
Jersey,
from
which
in California.
performed technical
Tommy
control
mosquito
Station,
which
commissions.
He
assisted
in the
mosquito
state' s
surveillance activities,
familiarize him
with
the problems.
This failed to
NUMBER 2
family
with
DECEMBER, 1993
household
goods to
stay. He
California, in
that
New
Jersey
could
admitted to
New
Jersey
within the
fit
in 1977.
his
retirement as
Specialist in 1975.
In
order
utilize
his broad
programs
technical
services
VOLUME 18
of
local
passed
away
on
12 August 1993, he
was
85.
He is
technicians
Richard F. Peters
Environmental Protection
Retired Chief
career
VOLUME 18
DECEMBER, 1993
NUMBER 2
IN MEMORIAM
John M. Doll
1935 - 1993,,
40
4441
off
4111100,
We
passed
John
away
University
He
Sunday,
November 21,
grew
science
army
on
in 1957.
He
went
on
to earn
in the
as a research scientist at
a masters
degree in
medical
In 1968, John
and
his
family
moved to
Syracuse, New
sorely missed.
Craig E. Levy
John
went
back to
microbiology in 1974,
at the
his doctorate in
University
of
Arizona.
xi
DECEMBER, 1993
VOLUME 18
NUMBER 2
IN MEMORIAM
Tuti R. Hadi
1943 - 1993
Ihor
Indonesia
irreplaceable
Hadi
and
the
asset on
jaw
of the
rest
world
13 March 1993,
when
Tuti Hadi
lost
an
mammalogistsanddevelopedcollaborativestudieswith
Dr. Tuti
born
although
con-
the
on
succumbed to meningitis.
25 July 1943, in
ducted her
life
research
Bandung,
centered
Java,
around
throughout the
of
degrees,
was completed
a second masters
Hawaii
at
American
Java,
she
wide span of
including
and doctoral(
1989)
in Jakarta,
in Javanese
Magelang, orYogyakarta.
and
her training,
was
degree ( 1980)
schools
University
of
many
at the
Xii
DECEMBER, 1993
NUMBER 2
reviewed articles.
expertise
is
The breadth
reflected
of
in the topics
her interests
of some of
Lance A. Durden
and
these
Statesboro, GA 30460
especially
ectoparasites),
VOLUME 18
the effect of
most
detailed
works pertained
Southeast Asia)
vectors.
and to the
nesian scientific
in many
taxonomy
of
of
Guy G. Musser
Department of Mammalogy,
American Museum of Natural History,
to scrub
parts
of
the chigger
major source
of her important
We
are grateful to
Pat W. Carney
work.
of the
on
Arlington, VA 22209
Dr. Hadi.
xiii
DECEMBER, 1993
BOOK REVIEW
A. R.
and
important animals.
Importance: A
Veterinary
Allied Terms'
microorganisms,
Pittaway
arthropod- transmitted
viruses,
1991
Distributor:
meticulous about
and
literature,
other
time or another
one
at
which
their selected
very
the proper
parasitology,
papers
textbooks
and
dealing
nonentomological
veterinary,
occasionally may include
which
etc.)
for
( medical,
it is
these
importance.
difficult
sometimes
and
A remedy
single,
has been
in
may
order to
finally
to these problems
written
that
arise,
is
specifically
One
available.
addresses
and
Allied Terms
source
was compiled
as a reliable
Society
of
America).
position of
to serve
of arthropods worldwide
in the fields
veterinary entomology.
of medical and
the author).
be
of
and reliable
information regarding
Also included
communicated.
biological
control agents(
employed
to
manage
intermediate hosts,
including
these
certain
diseases in
selected
lists
arthropod
natural
organismscarriedbythesemedical/
nomenclature to
are
vectors,
enemies,
and
veterinary arthropods
man
and
economically
saddleback caterpillar(
Io
moth ( Automeris
garden spider(
Sibine
stimulea[
Clemeus]), the
Argiope
trifasciata [ Forskal]),
the bird
Reviewed by D. L.Baumgartner, William Rainey Harper College, Department of Biological Sciences, 1200 West
Algonquin Road, Palatine, Illinois 60067, U. S. A.
76
Lytta
vesicatoria [
cinerea [ Forster]),
Linn.]),
and
Spanish
fly
any Neotropical
is
these few
otherwise
commendable.
Judging
from its
contents, the
primary
audience of
doctors,
veterinarians,
DECEMBER, 1993
have
limited knowledge
would
not
single
common
name
is listed.
Additional
but
of arthropods
who
frequently
read
by
inaccuracies. To these
would
be
an
orsimply propagate
invaluable reference.
of archaic and
may
incorrectly
older
literature
taxon.
limited
currently
Otherwise,
this reference
usefulness to
entomologists.
The
practicing
book
in that particular
would
medical and
be
veterinary
to
to
of only
occasionally" dabble"
with
medical/ veterinary
DECEMBER, 1993
ABSTRACT: Lyme disease has been reported from Australia based on clinical and serological diagnoses, but a
causative organism has not been identified. Attempts were made to detect, isolate, and identify spirochaetes from
ticks collected in areas associated with putative human infections. Ticks were collected by dragging in natural
bushland and by removal from native and domestic animals. They were dissected and a portion of the gut was
examined for spirochaetes by microscopy, with the remainder inoculated into culture media. The cultures were
examined for spirochaetes by microscopy and with immunochemical and molecular( PCR) techniques. Whole ticks
were also tested with PCR for spirochaetes. Between 1990 and 1992, more than 20,000 ticks were collected and
approximately 12, 000 were processed for spirochaetes. No evidence of Borrelia burgdorferi or other spirochaete
was detected. Some spirochaete-like objects detected in culture were shown to be artifacts, probably aggregates of
bacterial flagellae. The ecology of Ixodes holocyclus, the tick most likely to be involved as a vector to humans is
discussed in the context of a possible local spirochaetosis resembling Lyme disease.
INTRODUCTION
unpublished
data).
from North
and also
reported
Sigal 1988).
doubt
as
to the
hemisphere in
southern
The first
particular.
in Australia
reported
Japan
remains considerable
true
existence of
and
in Australia in
and
general,
disease'
a reference antigen.
South Wales
Hunter
Valley
region of
Further clinical
coast (
et al.
cases were
McCrossin 1986)
1986)
In
of
1988,
Lawrence
serological
was
diagnostic
service
for
IFAT)
and
through
tested.
tests to
1992,
specimens
indicate
2. 7%)
Lyme disease,
patients
to the
CDC
seven
and
clinical
( D. Dickeson
and
Although
in both
less than 1
for
criteria
patients
were
Lyme disease
Lyme disease
including
Australia
from 2, 446
possible
percent conformed
outside
suspected clinical
Only 66(
strain ( B31) of
referred with
This would
at
American
1988
enzyme
Westmead Hospital in
initiated
serological
the state.
a
Lyme disease
infected
R. Munro,
update
with
the
conclusions
of
the
three- year
investigation.
Presented at the 7th European Annual Meeting of SOVE, Bologna, Italy, August 27, 1992.
2Department of Medical Entomology, University ofS ydney, Westmead Hospital, Westmead, NSW 2145, Australia.
3Department
of
Microbiology,
78
DECEMBER, 1993
Tick
in the
by ` flagging'
were collected
removal
the
coastal
and `
border in the
identified,
dragging,'
animals
in
and
by
from
areas
Victorian
south.
Isolation
of
spirochaetes
The ticks
attempted.
N. S. W. Ticks
of
strip
Queensland border in
live
was
until processed,
in 70
by dipping
surface sterilized
twice in distilled
ethanol, washed
were stored
water, and
percent
the midgut
objects (
tissues dissected out aseptically. A portion of the midgut contents was examined
inoculated
into BSKII
1984)
incubated
culture media (
at
Johnson
33 C for up
et al.
and
were sought
by
Molecular
objects (
selective
in
isolated from
cultures
conditions.
cultures.
was
isolates'
attempted with a
including
characterization of
SLOs) found in
and
Media
to three months.
SLOs).
four polyclonal
Eighteen
such
and three
the OspA gene were obtained from just one` isolate' out
The mono-
clonals were
OspA
and
used to
directed
OspB,
and
flagellin
detect Borrelia
proteins.
specific
The PCR
DNA in
was
purified and
of B.
were
ribosomal gene(
burgdorferi(
a North
experiment
in
controls
were
also run
in
each
order
yielded no products
specific to
B. burgdorferi.
amplification products.
used to examine
American isolate)
41kDa flagellin
gene(
Persing
et al.
1990).
January
1990
and
were
collected
and
study
region were
DECEMBER, 1993
79
TABLE 1. Numbers of ticks collected for spirochaete isolation, January 1990- December 1992.
Species
Larvae
Nymphs
Males
Females
TOTAL
Amblyomma
moreliae
16
41
12
69
A. triguttatum
17
1633
513
79
148
2373
346
2176
66
1681
4269
7420
2057
561
2117
12155
66
46
10
77
199
175
68
245
1033
1033
22
31
10677
4880
784
4062
20403
Aponomma
concolor
Haemaphysalis
bancrofti
H. longicornis
Ixodes
cornuatus
I.fecialis
I. holocyclus
I. tasmani
1. trichosuri
Ixodes
species
Rhipicephalus
sanguineus
TOTAL
Borrelia.
In
contrast,
controls
relegated to
Oliver
et al.
DISCUSSION
disease
outside
known
and controversial.
of
endemic areas
is
of Lyme
problematical
infection( i. e. isolation
vectors, vertebrate
hosts,
of causative organism
or
humans),
positive
from
serology
80
DECEMBER, 1993
TABLE 2. Total number of ticks dissected for spirochaete isolation, January 1990- December 1992.
Species
Larvae
Nymphs
Males
Females
TOTAL
Amblyomma
0
14
554
265
67
71
957
232
914
18
748
1912
4862
1115
411
964
7352
I. tasmani
41
29
31
105
I. trichosuri
13
46
60
529
529
18
25
6231
2379
528
1832
10970
moreliae
A. triguttatum
Aponomma
concolor
Haemaphysalis
bancrofti
H. longicornis
Ixodes
cornuatus
I.fecialis
I. holocyclus
Ixodes
species
Rhipicephalus
sanguineus
TOTAL
notwithstanding
ultimately
spirochaetes.
The
these Australian
of similar
by
shown
microscopy
were
be
not to
SLOs
from
in
consisted of aggregations of
recovered
the SLOs
concession,
electron
cultures of ticks
flagellae thought to
originate
from
Australia.
Miles,
group
Similar
in
elsewhere
to
in Australia
B. burgdorferi
and
from
workers
pers. comm.).
objects
cultures
be
related
species
81
DECEMBER, 1993
4.
fir
r4.,
tl"
JO/
461.
Figure 1.
t.
Dark field micrograph of spirochaete-like objects( SLOs) detected in cultures from ticks, showing the
typically rigid and straight appearance in various lengths, and the uniform coiling( x400).
sc
et,.
f
j'
1` {
4.
s
A
t
k
gyp
',
Zit= ,
a
liti,
r'tdpitt.
Figure 2.
r,
Electron micrograph of portion of one of the spirochaete- like objects ( SLOs) in cultures from ticks,
showing
an apparent composition of
fiber-like
subunits( x5000).
82
DECEMBER, 1993
TABLE 3. Number of ticks examined by PCR for the presence of Borrelia, January 1990- December 1992.
Species
Larvae
Nymphs
Males
Females
TOTAL
Amblyomma
0
119
135
159
137
296
I.fecialis
I. holocyclus
279
53
236
570
I. tasmani
13
19
I. trichosuri
13
13
Ixodes
576
62
396
1038
moreliae
A. triguttatum
Aponomma
concolor
Haemaphysalis
bancrofti
H. longicornis
Ixodes
cornuatus
species
Rhipicephalus
sanguineus
TOTAL
However
tick in
Australia, belongs to
ricinuscomplex. Ixodes
the
holocyclus is unable to
B. burgdorferi
or
with
possible
Australian
strain of
spirochaetes
maintain
remains
unresolved.
Any
consideration
B. burgdorferi
should take
into
or a
of
similar
account
reservoir
spirochaete
hosts
for
in Australia
to that of
northern
identified
However, if there is
as reservoir
local
DECEMBER, 1993
B. burgdorferi transmitted
Lyme disease- like
by
in Australia, it has
83
Ramaciotti Foundations.
not
might
be
considered
syndrome
to be uncommon or rare.
appears
follows that
of
REFERENCES CITED
important
stage
northern
less
soon
noticeable
after
and
less
In
attachment.
Alpers, J.
1992.
be
southeastern
likely
to
1610- 1616.
of possible
be
during
infection) in
it is
when
the nymphs
human
in tick infested
likely
it is the larval
least
ovarially) that
areas,
be
Australia.
stages (
increasing
in
or near
residential areas
page A30.
Proc. V
disease, aproven
from
causative organism
has
not
vertebrate
host
cycles
in
southeastern
Australia
are
uncommon,
if not rare,
phenomenon.
The investigation
grant
was
Council
of
Australia,
and
a research
Medical Research
supported
and
by
from the
84
1991.
Russell, R. C.
situation
in
DECEMBER, 1993
Sigal, L. H.
1988.
borreliosis. Clin.
Sigal, L. H.
and
Exp.
Sinsky, R. J.
and
punch
biopsy
275.
DECEMBER, 1993
M. W. Service2
ABSTRACT: The main issue of this paper is whether we need more ecological data for developing control strategies
aimed at vector-borne diseases. Some argue that we already have sufficient knowledge of vectors, but need more
and better trained vector control specialists. In contrast, others believe our failure to control many vector-borne
diseases is due to insufficient understanding of vector ecology. With the prospect of decreasing budgets the question
is, should research be concentrated on topics that appear relevant to control, or is there still room for academic
may have" spin- offs" relevant to control? With some diseases, like dengue, we may already have
sufficient information on the vectors to decide on the best control approach, which in this case is source reduction.
research that
With other diseases, such as malaria and leishmaniasis more research on the vectors is needed to identify the most
appropriate control methods. Mathematical models concerning vector-borne disease control date back to Ross' s
1911) malaria model, but have they proved useful in helping design better control operations? A problem seems to
be that models have often promised more than they have delivered, and as a consequence models tend to be regarded
with suspicion by field entomologists. However, more recently, by unraveling the complexities of species
interactions, some models have provided valuable insights into population ecology of vectors and appear to have
been successful in identifying the best approaches for control. Nevertheless, despite the proliferation of models, it
must be admitted that as yet they have not often led to the successful implementation of better control. However,
despite these criticisms, I believe modelers and ecologists should be encouraged to work together to try and build
more realistic models that help both vector control specialists and epidemiologists.
relation to their behavior and habitats, and iv) the
WHAT IS ECOLOGY?
but I
It has
describe
be easy to
was mistaken.
let
big
can
hope to do is identify
proved
its
role, past or
future, in
subject,
vector control.
my
own
mostly
interests.
Firstly,
what
Ecology
is too
alone
on mosquitoes, that
data in
probably
control
reflect
is ecology,
apart that
is from
having
modeling.
Colinaux 1973).
been described
have been
community ecology,
ecology,
parasite
ecology,
mathematical
Ecology has
scientific natural history,
ecology.
1Paper presented at the 1st International Congress of Vector Ecology, San Diego, California, USA, 4 October 1993.
2Liverpool School of Tropical Medicine, Vector Biology and Control, Pembroke Place, Liverpool L3 5QA,
ENGLAND.
86
in the
organisms, as reflected
of diseases
in host
distribution
heterogeneity
non- random
populations, as well as
be
systems
reporting
frequently
extend
is
studies. Then, however, because of the lack of precontrol biological data, it is difficult to decide whether
size
demographic data.
interest,
and
other
Nevertheless, despite
and
ecology
little from
envied
by
ecologists,
over
strength
that
such
similarities
of
cross
DECEMBER, 1993
fertilization.
modern
biological
simple
only
chemical
limited
based on environmental
few
methods,
Such
agents.
amount of
virtually
pesticides
manipulation,
were
and use of a
approaches
required
biological knowledge,
such as
Commission
aspects"(
the need
was
based
on a comprehensive
unnecessary luxury.
declaration in a 1960 issue
an
by
is
eradication
above
administrative problem,
need to
of
World Health
published
the
know
about
its
all
exophily,
and
inadequate,
and
almost all we
survival
and this
organizational
an
rates
forced
of
vectors
was
often
Training
priorities
behaviour,
despite
some
and
and control" (
past
health
lessons there is
still a
lack
of
interest
by
strings
on
argued for more research, pointing out that markrecapture techniques had been invaluable in elucidating
many relevant aspects of vector behavior. Saying that
Furthermore,
as
long
as
control
remains
effective,
Malaria
control campaigns
in the
past, such as
DECEMBER, 1993
during
the
building
development
limited
ecological
epidemiological
sufficient
of
the Tennessee
of
Valley,
information
succeeded with
because there
largely
models,
87
biological information
was
It is
acknowledged
level management,
water
principally
were
based
good
in Malaysian
Italy
marshes of
Assam
and controlled
immediate
and
It is
dosing
and
including
streams with
forces
in the
protecting
communities.
onchocerciasis,
the work
key issues. Weknow that immigration and strong densitydependence can make any sterile male release program
biology
vector
of
African
where
DDT
the
of
about
because there
possible
blindness
were
fragile
Consequently,
was
there
In contrast, ecological
data have
ecological
Success
discrete foci
niche and
of river
nevertheless eradicated
indispensable in the
operation of the
For example,
proved
complex continued to
effective
larvicidal
very
coming from?
Careful biochemical
biting despite
caught
control.
therefore
Simulium damnosum
of the
sirbanum(
Townson
and
circumstantial evidence
outside the
OCP
transported
Meredith 1979),
strong
in the
area
while
being
on
as much as
damnosum
could
500 km. We
fly long
always
knew that S.
extent of
failure to
understand the
behavior
of a
vector
in the OCP
area,
non-
life( Yameogo
et al.
target organisms,
there is
Awareness
by
differences
in
their
degree
of
to
some
their
vertical
stratification (
theirimportanceinregulating
S. damnosum
of the need
Petersen
even now
of
apparently
1988). Nevertheless
including
aquatic
predators,
are
complex.
for
ecological studies
1983),
is
working in
understanding of the
ecology of the aquatic stages of Simulium quadrivittatum
provided
et al. (
a thorough
who
DECEMBER, 1993
88
to
was the
problem- solving.
prey.
It may be easy to find breeding places of mosquitoes but prove difficult to devise a sampling program
predators and
opposed to"
in terms
identify
that will
adults.
insecticides
density- independent
catastrophic
cause
processes.
biological
control
fashionable to
now
this usually
and
advocate
necessitates
an
introduce
aquatic
processes.
control
life
late
to kill the
processes
compensate
many
data
of
wealth
not
accumulation of research
control.
sad state of
in
by
DeFoliart
This
1987),
et al. (
decades
have
we
profited
little
transmission through
ecologists collected
have
or
control
with regard
vector control.
data that is
workers
last few
biologists
correct.
it
is this?
to
ecological
Molecular
remarkable
answers to these
their turn to
of opportunities to
simple questions,
apparently
dazzle, if not blind
us, with
find the
it is
now
biotechnology
and mathematics.
Should
we
continue
to reducing
Why
ignored
are
who
not relevant
studies?
that we
and
In fact, there is
despite the
control,
Have
are important in
led to
ecological
1970s has
since the
viral
Ro)
increasing
discussing
rate (
The
reproduction
mortality
epidemiological
stages of
reduces
mortality
Critical Thresholds
and
know
ecological
we can never
Kay ( 1985)
mean
DECEMBER, 1993
1990).
rise to
89
disease
significant
et al.
when the
Paradoxically, when
in both WEE
and
Reeves
processes(
et a1.
attack rates
switch
Olson
infection
and
1979)
et al.(
human
areas
many
mosquito
studied
California
of
SLE
and
correlation
over
21
reported
of all
from
years
dengue cases.
Culicineandanopheline- transmittedfilariasisdiffer
a positive
three factors.
peaked when
for SLE.
to ten
to
to
factor
in
urban areas.
Dye ( 1992)
to culicine- vectored
filariasis
and "
facilitation"
interpret the
in
to
traps placed
relationships.
rural areas
1979).
were
so
Despite
densities
cannot
say
densities
some
success
Cx. tarsalis
of
with
any
with
correlating
critical
be
to prevent viral
when as
factors
vectors
long
must
ago as
influencing
less information
reduced
the
of
ecology
some
WEE. We have
50
even
which
does
not occur
in
culicines.
Webber and
Culex
sampling
insensitive. However, Culex
ovitraps as developed by Reiter( 1987)
may help provide
on the population
threshold of
a reliable
density
have
proved
sampling
for establishing
method
thresholds.
find
no
significant
numbers of adult
any
of the three
Ae.
in
from houses
and
ovitraps.
Similarly
reported
in Honduras,
and
in
adult
is epidemiologically important
Despite
in evaluating the
transmission
has
because there
measuring
never
effectiveness
all
of
et al. (
numbers caught
serious
1990)
aegypti caught
Gil-Bellorin ( 1991)
association
larval indices,
of eggs collected
critical
densities.
Moreover,
much
DECEMBER, 1993
90
incidence
an
produce
transmission
of new
Therefore,
microfilaraemia.
intensity
be
must
in
cases
Wuchereria
know
Dye( 1992)
greater
with
Onchocerca
by
West Africa
level
exceeding 100
yearly( Annual
ATP), has been arbitrarily
deposition
volvulus
the
to
needed
Annual
larvae
be
and
not
combined
will
Brugia
density
critical
of
analysis
dynamics
population
whether there
A
bites
critical
more
efficiency
incidence rate of new
of
believes
for measuring
and improved,
refined
level
detectable
with
methods
not
on a person
of control of river
for
blindness to be
an acceptable
attained.
These
be prohibitive.
sufficient to
define
an
blindness
can still
The
project area
made
for any
maintained
However,
some
now covers
General Considerations
river
years.
be
by
ABR
at
be allowed only
should
level
after the
even with a
of transmission.
geographical variations
and
in biting intensities
sanitary
policy
of
mosquito
reduction"
to
the
capacity
itself( De Zoysa
in
immigration
of infected vectors or
improbable
situation
below
1991; Molineaux
et al.
vectorial
the practical
and
importance
This is because it
of
biting density
relies on an
densities
be
required
in
exists,
areas of
intense transmission
can
as
has been
over
parts of a
60
studied
in the Kisumu
area of
Kenya for
years,
geographic
areas,
developing
measures
to
tailored to specific
consideration
that are
be
must
given
epidemiological
differences
shifted
DECEMBER, 1993
91
Cox 1992).
hyperendemic
Nigeria, about
Modeling
in the Garki
area, such as
area of northern
1991).
This
is virtually impossible to
May
achieve,
late
It
when we recall
especially
a
cheap
can
aim
be
however, that it is
argued,
develop
vaccine
that is
immunity
partial
giving
more pragmatic to
northern
malaria control
Nigeria,
computer
Molineaux
on
instance,
of
basic
chemotherapy
of malaria epidemiology
Gramiccia 1980). In describing the
the project, Najera ( 1974) put the
of
discrepancies between
reproduction
rate
of
is
were,
for
estimates of
the
derived from
malaria
data. This is
often caused
getting
Macdonald'
on
of
and
the aid of
s
the absence
wide
model
and
shortcomings
blame
based
in
research project
simulations
mathematical
field
employed mass
and pregnant
children
The Garki
not an
by difficulties
seemingly
simple
found that
of
75
publications (
1974 to 1978) on
in host
heterogeneity
epidemiological
models.
Hasibeder 1986) to
by
the
selection
They
by
Dye
in
earlier.
and
predictions
wondered (
Garki
vectors
Molineaux
and
generated
biting
malaria model(
ignored. Such
heterogeneity
in behavior appears to be common in biology. A readable
of non- random
was
account of mathematical
medically important
last 20
modelers,
population
Reeves
et al. (
ecology
and control of
Cx. tarsalis.
For
increasingly
populations.
occupy discrete habitats such as containerbreeding mosquitoes because their biology lends them
vector efficiency.
Despite all the above criticisms and constraints, I
years,
and
animals that
to
having
several
generations.
Many
of the
important
vector
heterogeneous
92
biology
aspects of a vector' s
Simulation
have clearly
identified adult vector survival rates as the most important
single
factor affecting
Reducing
for
experiments,
survival
example,
infection
population
rates
rates.
logarithmically
can
1991)
reduce
gives
very
DECEMBER, 1993
temperatures.
In Louisiana, Focks
et al. (
and although
many
of the points
he
interest.
of
Modeling
Pondicherry, including
Das ( 1987), built
deterministic
models
for
general,
further evaluation.
her
in
data,
example, the
infected
and
simple
filariasis transmission. In
the dynamics of
except
for
mosquito
infection
rates.
For
in
mosquitoes
underestimated
Various
an
area
it in
under
explanations
discrepancies,
lacking
an area
including
but
vector control.
control
vector
were
for
offered
heterogeneity
possible
these
in the
model
for
used
adult
emergence, such as
would
be the
by source
infection,
is
which
in filariasis
for
larviciding,
interrupting
mean of some
Because
reduction or
that a
was required to
similar to previous
studies.
Cx. tarsalis in
been
the
of
importance
several attempts to
predict population
strategies(
develop
of
computer models
debatable
whether
any
effective
control.
Greever
model
has
and
resulted
in
more
Georghiou ( 1979)
For
instance, it did
vectors, and
data
not take
they
into
they had
inadequacy of the
available on which
They
to
invited
Georghiou ( person.
others to
comm.
improve
1993)
their
admits that
in California
appear
80 F
accumulation of 50
degree days
over
insecticidal
DECEMBER, 1993
have been
93
the 1960s,
methods which
advocated since
will
spraying
have little
likely
will
long- term
effect
on
adult
field
at
workers
It is
way to
control
dengue is
only truly
source reduction."
interesting
key meteorological
inappropriate,
so
his
conclusions were
wrong!
More
factor governing
temperature as the
Portier
1985). In
et al.
found to be
useful
rainfall was,
based
on rainfall
data
Birley
and
decisions. He
be less costly
provide
a means of
control
making
larval
biggest
world' s
far
has been
put
into this
surveys.
surpasses that
research than
not
by
predict epidemiological
1992, Plaisier
1986) built
et al.
1990). For
example,
a catalytic model to
epidemiological trends
during
Remme
et al.
age- specific
study
spraying intervals
Why? It is
undoubtedly because of logistic problems. A weekly
would
save
money.
repeated on
different days.
Also,
Instead, they
index
was
years
Community
to
widely
Further modeling, 12 to 14
initiated, indicated that because
or more, which
and
is
as the
now
used
of the
Remme
1990).
They
found
a good
observed
epidemiologicalandentomologicaldata collectedfrom
vector
planned(
agreement
epidemiological
during
were
et al.
between
predicted
some
also
and
Nevertheless there
discrepancies between
predictions
and
and
chemotherapeutic
control
strategies,
94
damnosum
volvulus would
at six- month
be
years could
if ivermectin
contrast,
intervals, it
in 18
eradicated
before 14
control
In
recrudescence.
delivered
to
O.
but abandoning
lead to
years
the village.
entering
99
with
SIMON
or people
that
predicted
were
up to 29
feasible
could take
stop
levels,
be 99
self-sustaining,
both
drug
control,
disadvantage
trol, although a
at reduced
vector and
could
residual
because the
longer be
Combined
DECEMBER, 1993
of
from the
elimination of worms
population.
The
model
a greater
these
relevant
from the
consequently be
of
people
data
epidemiological
were
collected
actually
CONCLUSIONS
There
further
improving
Ae.
can
be few
generalizations on
For
control strategies.
albopictus are
including
example,
Ae.
for
aegypti and
viruses, and
both
in
occur
are
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Similarly it is generally
1991.
Anderson, R. M.
most appropriate
dengue
60: 1- 50.
outbreak.
areas (
Service 1992)
result of variations
rates, and not
for
in house
and appears to
construction and
ecological reasons.
So it
be the
dosage
be
might
Anderson, R. M.
and
R. M.
May.
1991. Infectious
control.
Anopheles
In
species
marked contrast,
transmitting
malaria
and
and
for
vector
their
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spraying houses
exophilic.
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insecticides if the
A complication is
use
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is
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1927.
1981.
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very
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Aedes
Dominico,
Pen
DECEMBER, 1993
ABSTRACT: Adult and immature stages of Ixodes scapularis Say( previously Ixodes dammini Spielman, Clifford,
Piesman, and Corwin) from a study area in Kent County, Maryland, were tested for infection with Borrelia
burgdorferi by immunofluorescence assay in 1991- 1992. Infection rates ranged from 27.0 percent to 45.2 percent
for different life stages, sexes, generations, and seasons of activity. The infection rates of engorged larvae attached
to white- footed mice( Peromyscus leucopus[ Rafinesque]) and the succeeding generation of questing nymphs were
similar, indicating that mice were the chief spirochete reservoir hosts in the area. Infection rates were significantly
higher in questing adults than in their nymphal progenitors, indicating that the tick population acquired a substantial
portion of infection during nymphal feeding. Estimates of the intensity of spirochete infection per tick( spirochete
burden) showed significant( P< 0.05) increases in spirochete numbers from larva to adult but not between nymphs
of
different
generations or adults of
different
seasons.
overwintering infected adult ticks. A significant, sex- related difference in spirochete burdens was seen in adults.
Studies from other areas of the I.scapularis geographic range would help to clarify seasonal and sex-related trends
in infection rates and spirochete burdens observed to date.
INTRODUCTION
Ixodes
Say ( reported
scapularis
Corwin[ Oliver et
Lyme disease in
al.
the junior
the northeastern
virginianus
season
reported as I.dammini)
have
Zimmerman)
shown
with
the
infected
during
that I. scapularis
Maryland, and is
spirochete Borrelia
Steigerwalt,
may be
increasing
Amerasinghe
et al.
1993).
life
exists on
the ecology of
County, Maryland,
annual temperature=
[ NOAA]).
Pinus taeda).
that
showed
winter months
We
( March- May)
of
1992.
of
Entomology, University
of
other
white- footed
from
collected
mice (
P. leucopus)
during
mammals
small
and
the
summer(
set out at
area.
in the study
United States (
eastern
Haught 1983),
area(
and also
in Lane
reviewed
Ticks
halothane
recovery.
kept within humidified vials at4 C until
were
identification( based
on
spirochete
Midgut
is
1991).
et al.
smears of
et al.
1993)
days
within seven
1991
and
and
n=
of 1992.
tested for
of
direct immuno-
the presence
of
B. burgdorferi
of collection.
were
per host.
processing for
and
LeConte]).
B. burgdorferi in the
reservoir of
shown
scapularis
The
DECEMBER, 1993
100
by
antibodies(
in three
et al.
1992).
fields
by counting
all spirochetes
320x
magnification
microscope
worker (
three
The
1-PA).
fields
infection(
of view
under
burden)
spirochete
vs.
nymphs vs.
Two- way
nymphs).
1992:
comparisons of spirochete
burdens in
is
previous
published
et al.
text
Fall-
summer( X2=
spring
ANOVA) on
adult ticks.
junior
when
and
spring 1992:
the name
under
relegated to a
I. scapularis( Oliver
2. 87, df= 1, P=
and
summer vs.
for
was used
X2=
infected
burdens( larvae
analysis of variance(
data
loge- transformed
adults,
Infection
for
nymphs,
excluded.
used
were
be
of
per tick.
intensity
Contingency
dammini.
X2=
X2=
of
168 larval
was collected on
mean
of
SD= 3. 28
from 797
1991.
37
trap- nights
Data
on
29
nymphal
I. scapularis
U Test,
per mouse)
in the
summer of
of collection
I. scapularis from
mammals were as
tailed shrews(
and
other
follows: 10 larvae
Blarina brevicaudata[
on
trapped
Say]);
of
short-
5. 13, df = 1,
P<
1991
nymphs ( z =
3. 11, df=
1, P = 0.002).
small
1 larva and
z =
z=
DECEMBER, 1993
101
TABLE 1. Borrelia burgdorferi infection rates in questing adult and nymphal Ixodes scapularis.
Percent Infected( No. tested)
Total
Males
Females
Adults
Summer 1991
27. 0( 74)
Fall 1991
1992
Spring
TABLE 2. Mean
45. 2( 62)
39. 4( 66)
42.2( 128)
41. 8( 55)
42.9( 70)
42.4( 125)
42.2( 64)
Host- Attached
Questing
Questing
Questing
Total
Larvaea
Nymphsb
Males
Femalesc
Adults
75. 3 50. 8
100. 5 58. 0
87. 9 54.6
51. 9 29. 7
96. 2 53. 4
74. 1
Summer 1991
6. 57 6. 36
37. 1
31. 1
Fall 1991
1992
Spring
Nymphs
57. 1 48. 4
47. 8
aBased on 20 larvae.
Note: Spirochete burden was significantly less in males than females ( fall 1991 and spring 1992 adults, 2- way
ANOVA, P= 0. 04).
spring 1992( Mann- Whitney U test, P< 0. 001). All other comparisons within adult or nymphal cohorts were
Statistical
burden in
spirochete
ANOVA)
of
the
spring 1992)
non- significant(
spirochete
P=
0.41
and
significantly
DISCUSSION
seasonally
change
spirochete
of nymphal
I.
scapularis, or
between
during
during
DECEMBER, 1993
102
of
2).
be
successive
larval and
feeding cycles.
have been
Rates
shown
to
was seen
in Lane
in
infection
peak
adult ticks.
this is important in
in
in
rates
questionable whether
spirochete maintenance
of adult
of
seasonality
et al.
their
I. scapularis in the
not
mammals
like deer,
skunks,
raccoons,
and
opossums
not
hosts.
and
spirochetes to
skunks
I.
of
lower
capable
are
scapularis
but
at
that raccoons
transmitting
1990), both
1985).
during
nymphs
disease epidemiology or in
nature. The guesting period
shown
et al. (
feeding
1991, Spielman
et al.
period results
et al.
Piesman
spirochete transmission
of more
sized
of
to humans
in
as
The infection
northeastern
in the study
play
in nature as well
However, it is
et al.
succeeding nymphs
in the maintenance cycle of B. burgdorferi
reviewed
Piesman
would
akey role
repletion(
reservoir
larvae
nymphal
transmission
transovarial
e.,
contend
efficiency
incompetent
are
decrease
Telford
1988)
for B. burgdorferi
reservoirs
spirochete transmission
Oliver
et al.(
host, i.
by
serving
1992) have
and
as
may
dead-
that
hosts.
end
et
al. (
asserted
PCR)
of
assays;
they did
not
in the deer
or
In
study
approximately
those of
Sharon
were
success.
Sharon
et al
et al. (
spirochete
methodology in Wisconsin.
burdens in ticks increased
eight- fold
our
of
the ticks
succeeding questing
sex
in Wisconsin.
nymphs, and
nymphs and
numbers recorded
been
an underestimate of
acquired
had
not
conclusions
based
on
Studies from
resolving the
sexes.
geographic
scapularis
103
DECEMBER, 1993
issues
range
I.
be helpful in
would
1992)
by Sharon
raised
and
W. Burgdorfer.
1991.
Acknowledgments
We
thank
1991.
Funding
and
during the
provided by the
assistance
was
Nancy
Varuni
W. A. Chappell.
1984.
Parasitism by Ixodes
summer of
Maryland
This is Scientific
REFERENCES CITED
1992.
Distribution,
and
Sweeney.
Lyme disease
spirochete
Ixodidae)
density,
K.
and
on
white- tailed
deer in Maryland.
1991.
562- 568.
J.
James, B. S. McGuire,
and
T. W. Scott. 1993.
Increasing density
and
Antibodies to Borrelia
of
Ixodes
and
E. Howerth.
1992.
white- tailed
deer.
J.
Ent.( in press).
Anderson, J. F., L. A. Magnarelli, W. Burgdorfer,
and
mammals
Med.
Hyg.
from Connecticut.
Am. J.
Trop.
and
of
the Lyme disease spirochete in populations of whitetailed deer and white- footed mice.
Yale J. Biol.
and
southern
J.
Spielman.
disease spirochetes( Borrelia burgdorferi) in fieldcollected larval Ixodes dammini( Acari: Ixodidae).
role of medium-
burgdorferi
in
scapularis and
345.
Thesis,
University
of
Maryland.
vi+
49 pp.
Wiesfeld, A. W. Hightower,
and
C. V. Broome.
104
1985.
Surveillance
United States,
1982.
Lyme disease
in
the
of
1149.
Sharon, M. D., W. A.
DECEMBER, 1993
Blacksburg, 44 pp.
Rowley, M. G. Novak, and K. B.
of Virginia: No 13.
Ticks of Virginia( Acari: Metastigmata). Research
Entomology, College
of
Department
Agriculture
and
of
Life
DECEMBER, 1993
Dave D. Chadeel
ABSTRACT: The wing lengths ofAedes aegypti females collected as pupae from cans, basins, buckets, and drums
averaged 2.42 mm, 2.61 mm, 2. 61 mm, and 2. 65 mm, respectively. Females with the largest wing lengths were
collected from drums( F=93. 62, P< 0.001). The wing lengths of adults emerging from drums ( P< 0.001), buckets
P< 0.01), and basins( P< 0. 01) were significantly larger than those
emerging from cans( P= 0. 21). The wing lengths
ofparous host- seeking females collected indoors were larger( 2. 59 mm) than parous females collected outdoors( 2. 53
mm) but this difference was not significant. In contrast, the nulliparous females collected outdoors( 2.49 mm) were
significantly( P< 0.01) larger than the nullipars collected indoors( 2. 36 mm). However, both nulliparous and parous
host-seeking females collected outdoors were similar in size when compared( 2.49 vs. 2.53 mm). The data suggested
that small Ae. aegypti females exhibit reduced blood- feeding success indoors and possibly reduced survival when
compared to large adults.
INTRODUCTION
due to
breeding
mosquitoes
factors
numerous
such
vary markedly in
as
overcrowding
Mulla 1979), temperature ( Brust 1967), mutual
interference ( Terzain
and insufficient
the
larval
However, the
be
amount of variation
related to the
in
ephemeral
body size
sites. Consequently, individual size is an important part
of the life- history strategy of a species and is often
related to survivorship and fecundity ( Begon et al.
1986). Nasci( 1986) reported great variations in wing
length among emerging Aedes aegypti males and
females and concluded that small females may exhibit
reduced blood- feeding success and most
likely reduced
survival when compared with
purpose of this
large
during
The
stage.
size appears to
and
adults.
study
wing
length ofAe. aegypti adults emerging from four different
to
Ae.
size of
aegypti
of
jars lined
collector and
In the
kept separate
females
by
parous
x magnification,
Ae.
The
Detinova' s
or absence
presence
1962). Mosquito
for
and counted.
40
method,
to
Data
the
parous
according
Detinova
were analyzed
using
collected
outdoors (
P> 0. 01)
were
separately
( indoor or
an analysis of var-
females
of tracheolar skeins (
outdoc:).
when compared.
identified,
aegypti
by
location.
or
DECEMBER, 1993
106
DISCUSSION
containers
RESULTS
Wing
length
measurements of
Ae.
in
No
wing
detected
lengths
for each pre- adult was different in each tire which may
aegypti adults
lengths
are presented
in TABLE 1.
of male and
when compared.
were collected
female Ae.
Females
with
and
right
aegypti were
The
TABLE 1.
In contrast, size
Wing length of emerging Aedes aegypti collected from drums, buckets, basins, and
cans in the field.
Sex
Sample
Average
Emerging
Emerging
247
2. 05+ 0. 05
1. 96- 2. 15
269
2. 65+ 0. 09
2.43- 2. 88
Emerging
Emerging
353
2. 03+ 0. 01
1. 74- 2. 31
335
2. 61+ 0. 01
2. 15- 2. 99
Emerging
Emerging
273
2. 02+ 0. 04
1. 84- 2. 12
267
2. 61+ 0. 07
2. 30- 2. 78
193
1. 93+ 0. 06
1. 74- 1. 74
208
2. 42+ 0. 09
1. 96-2. 78
Drums
Male
Drums
Female
Buckets
Male
Buckets
Female
Basins
Male
Basins
Female
Cans
Male
Cans
Female
Emerging
Emerging
Range
DECEMBER, 1993
TABLE 2.
107
Sample
Host- seeking
n)
Parous
Parous
Nulliparous
100)
28
2. 59+ 0.06
2. 36+ 0. 05
105)
21
2. 53+ 0. 07
2. 49+ 0. 05
indoors)
Host- seeking
outdoors)
are
frequently
more
Milby
fecundity(
sierriensis (
females
parous (
Say)
Bock
Ludlow)
and
in this
and
parity
to enter houses
likely
to blood feed.
During
of
aegypti were
found,
ACKNOWLEDGMENTS
quality
and the
each container(
in
Stahler 1949).
sites,
however, individuals
were
larger in
field.
REFERENCES
size
successfully
suggested
distances, are
blood
Nasci ( 1986)
meals.
that small
short
Ecology:
Individuals,
Populations
and
et al.(
fed
on an optimal
diet
adult Ae.
were more
female Ae.
smaller
in
cans
had
temperatures.
993.
collected.
in size to the
These
parous
results suggest
houses
aegypti ( L.)
DECEMBER, 1993
108
Chadee, D. D.
of
table characteristics
selected
aegypti(
L.).
Haramis, L. D.
1983.
72.
and
and
P. S. Corbel.
diel
1987.
Seasonal
in the
patterns of oviposition
of the mosquito,
Aedes
aegypti(
L.)( Diptera:
Trop. Med.
Christophers, S. R.
1960. Aedes
aegypti (
L.). The
216 pp.
Fish, D. 1985. An analysis of adult size variation within
populations. Ecology of
Pp. 419-429 in Proceedings of a
Workshop( L. P. Lounibos, J. Rey, and J. H. Frank,
natural
mosquito
Mosquitoes.
eds.).
of
and
P. Barbosa. 1971.
four Aedes
standard technique.
and
F. J. Rohlf.
1980.
Biometry. The
Sokal, R. R.
density
on
some
laboratory
DECEMBER, 1993
This laboratory study analyzes average daily survivorship and adult life span of Eretmapodites
Abstract:
quinquevittatus, Aedes aegypti, Aedes albopictus, and Aedes bahamensis. Experimental populations of 25 adult
males and 25 adult females were observed, as well as experimental pair groups of 25 pair cages containing one male/
one female each. Life spans of blood fed and non-blood fed populations of Er. quinquevittatus were compared to
those ofAe. aegypti, Ae. albopictus, andAe. bahamensis. Average daily survivorship for males of Er.quinquevittatus
was
significantly
greater than
species
regardless
of
feeding
treatment.
Average daily
survivorship for females of Er. quinquevittatus was significantly greater than that of Ae. aegypti in the non-blood
fed treatment. Non-blood fed average daily survivorship for Ae. bahamensis was significantly greater than that of
the other two Aedes species, and not significantly different than that of non-blood fed females ofEr. quinquevittatus.
Survivorship forAe. aegypti was greater when males and females were blood fed, which conflicts with earlier reports.
Unexpectedly, survivorship and longevity of Er. quinquevittatus males was significantly greater than that of other
commonly studied mosquitoes.
INTRODUCTION
is easily
quinquevittatus
an excellentorganism
reared
Africa
studies
in the
and
Hartberg
arboviruses (
Er.
show that
laboratory and is
and Gerberg
forresearch( Hartberg
1971).
This study
for Er.
compared average
when
quinquevittatus
experimental populations or
analyzed
daily
adults
individual
feeding
species.
survivorship
held in
were
We
pairs.
on average
We
also
daily
compared adult
1985,
and
Hawley
1988),
Weyer 1965).
because Aedes
and
Edwards 1941,
and
These
Hartberg
et al.
1984, Liu
history
of
1Department
of
Baylor
were maintained at
25 C, 80 5 percent
the
completed
studies on
Biology,
1968).
Faircloth 1983).
et al.
and
Aedes albopictus
Eretmapodites
University,
110
200
ml of
DECEMBER, 1993
RESULTS
of
inside the
towel for
To
populations
from the
were obtained
at
white-
eggs.
receiving
establish
All
pair cages.
following
colonies maintained
strain of
and
bahamensis. The
cylindrical,
pint- size
eggs were
emergence
These
quinquevittatus,
cultured and
emerged(>_10/ cup).
Er.
adults
cups
where
Days lived by
these adults
daily survivorship
and
instar larvae
30
26
with>
cm x
250
ml
were
fed
30 eggs
6 cm),
cm),
beakers,
of
slurry
half-pint
or
for the
blood to
in large pans
mason
data.
pint- size
jars. Larvae
top
adults
adult
and
and
production
anautogenous
meal
was
Twelve hours
after
blood
subsided.
egg
given.
blood
Subsequently,
meals
P< 0.05).
were
provided
foreach adult.
for
the
were
group( 25
also calculated
for males
each
and
day.
Mean life
Survivorship for Ae. bahamensis males held with nonblood fed females was significantly greater than the
survivorship of Ae. aegypti and Ae. albopictus males
span was
and pair
group.
Effects of
density
on adult
2- way ANOVA
with
transformed average
size,
life
blood
span were
replication
daily
feeding,
using
survivorships.
determine
and
determined
to
Survivorship for
surviving
by
arcsine
Fischer
PLSD)
were
which comparisons
among
differed.
Alpha and power for the
significantly
statistical tests were 0. 05 and 0.80,
respectively.
species
P< 0. 001).
1 female
performed
species compared(
the population
mosquito
four
documented
male/
Survivorship of
blood fed Ae. aegypti females was significantly greater
egg
P< 0. 001).
were
anesthetized mice
females.
compared (
emergence cups
of
species
10/ cup).
cubes placed on
carbohydrate
provided
were reared
Progeny
while
transferred to cylindrical,
Sugar
were transferred
development
32 cm
which
Eggs hatched
Ae.
did
DECEMBER, 1993
TABLE 1.
Average
daily
survivorship(%)
111
aegypti, Aedes albopictus, and Aedes bahamensis held in populations( 25 males/ 25 females each) and
pairs (
male/
1 female
each- 25
pairs
in
group) in the
laboratory.
parentheses.
Avg. Daily
Male
Eretmapodites
quinquevittatus
Aedes
aegypti
Sur(%)
Female
Female
96.4
97. 8
37. 6( 31. 8)
96.6
97. 4
57. 4( 38. 1)
77. 0(40.4)
96. 3
95. 1
38. 5( 33. 9)
25. 9( 24.8)
96. 6
96. 3
30.6(37.0)
38. 7( 31. 8)
96.9
97. 7
60. 2( 24. 7)
57. 5( 35. 3)
94.9
96. 7
35. 3( 26. 7)
47. 0( 32.2)
95. 5
95. 7
48. 0(26. 3)
41. 4( 21. 2)
96.4
90. 5
40. 2( 34. 3)
12. 8( 8. 3)
95. 8
94.4
42. 8( 28.9)
22.2( 17.4)
96. 1
91. 9
24. 6( 26. 1)
14.7( 7. 5)
96. 8
96. 2
56. 7( 35. 1)
43. 0( 31. 1)
96. 2
85. 7
54. 0(27. 4)
61. 0( 10.5)
96. 5
95. 8
50. 0( 35. 6)
44.3( 25. 3)
96. 2
97. 0
47. 2( 24. 8)
49. 2( 33. 1)
96. 1
96. 0
42. 9( 28. 8)
59. 7( 30.9)
95. 7
95. 7
42.9( 28.4)
55. 9( 22. 5)
96. 6
97. 5
67. 2( 34. 7)
47. 5( 45. 2)
96.9
97. 5
51. 6( 40. 9)
53. 7( 40.2)
96. 8
96.9
62. 8( 37.9)
75. 7( 30.8)
97. 3
96.6
77. 6( 38. 1)
71. 6( 30.2)
83. 4
92. 1
8. 3( 3. 2)
19. 8( 9. 5)
85. 0
97. 5
8. 4( 4.2)
57. 2( 40.2)
91. 2
96.9
9. 7( 7. 0)
60.9( 39.4)
53. 1( 49. 7)
82. 6
86. 6
8. 1( 3. 4)
20.3( 4. 9)
77.6
88. 3
9. 0( 2.8)
17.4( 6.9)
77. 7
87. 0
9. 4( 2.9)
16.9( 5. 9)
albopictus
83. 8
97. 4
11. 9( 4. 8)
57.6(42. 7)
80. 9
96.7
11. 2( 4.6)
45.0(35. 8)
91. 0
96.8
15. 1( 6.6)
46.2( 41. 1)
90.2
96. 1
14. 8( 9. 3)
49.2( 26.7)
85. 3
95. 7
10. 2( 4. 9)
43. 7( 32. 5)
84. 1
95. 6
11. 2( 26. 7)
41. 0( 33. 7)
Aedes bahamensis
83. 8
97. 3
13. 7( 5. 3)
63. 6( 44.9)
90.0
96.9
20.4( 8. 4)
65. 3( 33. 1)
94. 1
97. 1
33. 0( 18. 6)
44.0(38. 2)
92. 3
96.2
19.8( 12. 3)
52.0(26.0)
95. 5
96.2
29. 4( 20.6)
61. 8( 24. 5)
92.6
96.2
20. 3( 11. 9)
61. 7( 24. 5)
Aedes
112
not
differ due to
feeding
DECEMBER, 1993
Edwards, F. W.
treatment.
1941.
Average
daily
quinquevittatus
the
other
did
not
Aedes females,
quinquevittatus survived
only do the
while
P. Zhao,
and
N. H. Cao.
1984.
Studies on the
Er.
males.
Not
longer
of
males
499 pp.
1971.
Ae. bahamensis.
of
albopictus and
survived as
Ae.
long
aegypti.
as
Females
females
of
Er.
ofAe.
bahamensis
quinquevittatus.
If
Gubler, D. J.
1970.
Comparison of reproductive
males and
females
survived
longer
the blood fed treatment, than when under the non- blood
fed treatment.
We have
no
explanation
for this
occurrence.
en
Ethiopie
partir
de
collectes
exot.
58: 574-
589.
Hylton, A. R.
1969.
147- 149.
J. Salaun,
and
L. Ferrara. 1969. A
new arbovirus
mosquitoes
chrysogaster.
human
new record-
Liu, Z. W., Y. J.
Zhang,
Population dynamics
and
of
Y. Z.
Aedes
Yang.
1985.
( Stegomyia)
Christophers, S. R.
1960.
Aedes
aegypti:
Its Life
English summary).
of chicks
to
DECEMBER, 1993
Semiliki forest
virus ( Kumba
Strain). Ann.
Trop.
113
of spondweni virus
report of two
in
and
Ethiopia
5.
Isolation
arthropod vectors.
877.
Aedes ( Howardina)
albonotatus (
Coquillet),
of viral
yellow
fever
strains
from
38: 873-
Org.
583- 592.
DECEMBER, 1993
J. D. Lange
AB STRACT: As part of vector-borne disease surveillance activities, fleas were collected from sylvatic and domestic
mammals at 46 different localities throughout San Diego County from April, 1991, through December, 1992.
Twenty- four flea species( 12, 121 specimens) were collected from 26 of the 32 host taxa( 2, 397 specimens) examined.
Ten of these species have not been reported previously in the literature as occurring in San Diego County. A review
of unpublished flea surveys of this county yielded two additional species, Orchopeas leucopus and Pulex irritans,
which were not found during the present survey. Thirteen of the 26 flea species occurring in San Diego County have
been implicated as disease vectors. Oropsylla montana, the primary plague vector in the western United States,
comprised 62 percent of the specimens collected during this survey.
INTRODUCTION
The importance
other
a number of
relationships
between these
vectors and
surveys
and
fleas
of
ecological
their sylvatic
of sylvatic mammals
have
Stewart
Schwan
et al.
and
and
Brennan 1957),
and
Rutledge
as
1969, Jameson
Kinney
1979),
and southern (
Nelson 1983)
et
al.
1951,
Augustson 1943,
Murray
1971).
Fleas
also
have been
collected
and
in
southern
1957)
et al.
1986). Flea
et al.
surveys of
urban areas of
60060).
et
1985),
Eskey
and
1943).
of short
restricted to a
of
mammals
San Francisco(
domestic
from
coexisting in interfacial
al.
central
fleas
County
in
survey
southern
contributes to the
on sylvatic and
and
California.
domestic
identifies fleas
control purposes
during
Consequently,
the
biological knowledge
rodents
and their
in San Diego
flea-borne disease
possible
outbreak.
hosts for
Camp
MATERIALS AND METHODS
Pendleton
for fleas.
All
As
Lyme disease
surveillance
Record Form.
1Vector Surveillance and Control Division, San Diego County Department of Health Services, San Diego, CA
92186, U. S. A.
115
DECEMBER, 1993
total of
identified from 26
species, was
during
following
representing 24
Sciurus
Eads
1985).
A casual
33 host taxa
squirrel,
found
on the
2, 396).
include
results
of the
Fleas
The
specimens,
were
species
et al.
for
unpublished records
griseus(
County.
This campground is
Ceratophyllid Fleas
New Mexico
and
west to
California
and
et al.
Nelson
for
elevations(
Lang
Evans
western
largest
dine
with
0.
s. sexdentatus
County
Park( 1, 575
record
the
and
while
and
San
were suggested to
specimens
0.
of
present
survey
This flea was
October
The majority
of
the
ranges (
Haddow
et al.
1983).
during
the
months
of
January.
western
states,
the deer
sporadic
mouse
flea,
distribution
with
Augustson 1943). It
1943).
An
June, 1982,
at
three
an unpublished
examined
in
specimens
during
plague
the
sexdentatus collected
southern
more common
through
In the
s.
were
be
and
western
southern
Schwan
occurring in
1992). These
m)(
Schwan
m)(
sexdentatus and
to have a
sexdentatus, appear
are
a principal
elevations.
subspecies of the
Diego
number of
the present
flea, Orchopeas
during
western
et al.
sylvatic
north- south
et al.
Wills 1991).
and
collected
records
Surveys in
in the
Oropsylla
wood rat
1947).
while
In
1986,
fleas
spp.
plague
1989),
Peromyscus spp.,
1943,
higher
at the
et al.
montana
and
Oropsylla
host
on this
1951),
et al.
San Diego
in March, 1961,
Julian( 1, 274
m) which
116
6)
sninosnw
DECEMBER, 1993
snyy
N
O
ci,
1) vuviui8.nn siydiapij
V
z
1) i Cuoyluv
0,
1)
suvou8lu
snau8
aviloq
s(
sninios
uiouioy,L
8
snoivao
vin vo snlo
oi
mm
thwoluoponyiiag
L) t3ai p 1 iouvs
to
r,
a)snpnvoquoi
spoiv8aw
o`
r,
e
a
L)
1 (
aiOJ
i ( oq snoswoad
oz(*
isuaydais snuuo snos( woad
as
E
N
c,
08)
iiiagwv8 snivinoiurnu
1'
601) sninalaivrfsnoiwaia
snos(
snosLCwo1ad
woaad
11
9)
su8su
snoiJofivo
snosCwoiad
c)
In
en
en
a
0
1(
0(,
O
'
i7
..
1 L.)
i
K =
ra
t7
r
N
v
O
C
8V
In
tZ
wi
_
i
1
y
2.
000
o
43
O
y
E-
C7
GL
Vg g
az
C7
Marchette
1962).
et al.
Peromyscus
During
M.
of
specimen
single
117
DECEMBER, 1993
from
collected
was
sinomus
in June,
The deer
mouse
into
wagneri ophidius,
and
Pulicid Fleas
forest.
flea, Aetheca
The
southern
abundance of
wagneri was
rarely being
Lava Beds National Monument, Siskiyou
northern
California
records are
Kinney
10
report
fraterculus)
1988). The
be
scarce
present
flea
were
with
Most( 11)
the
w. ophidius to
majority( 9
of
which
m)
San Francisco
also
ranges
from the
into Nevada
Bay
and
other
on
1988).
Observatory
predominated
by
collected
from a T. b.
Campground ( 1, 477
Mountain ( TABLE
1).
This
Mus
nigricans
Palomar
m),
is
Hubbard 1947).
occurs on
Rattus
on
other
occasionally
In Los Angeles County, N.
and
flea found
on
index( 2. 2) occurring
Schwan et al. 1985). In
on
Rattus
while an unpublished
1947).
musculus,
with
a male and
campground
Rattusrattus
Hubbard 1947).
fasciatus
2).
single specimen
and
1943), Kern
w. ophidius varies
A.
mixed chaparral
spp.
et al.
rodents (
in Alameda ( Meyer
et al.
Lewis
from
plague
1979).
with
et al.
is
locally, preferring
rare
1971,
specimens of this
These fmdings
The
Ryckman
in main-
County
found
was
1951,
Mataguay
at
a minor role
found A.
also
County,
m. gambelii.
characterized
flea
survey
eremicus
P.
in San Diego
13) from P.
may play
six
in June, 1982,
wagneri
in
plague
from
specimens
examined
Creek. Aetheca
taming
Orange Counties
and
unpublished
records
County,
1969). Southern
norvegicus
1987 survey
et
al.
1988).
In southern California, it
Although N.fasciatus
is
may cause some human plague cases when the index
maintain
or
to
initiate
epizootics
high, it was unlikely to
also
found it
each on
and on
common on
M. musculus( 1
S. beecheyi(
R.
examined),
nudipes)(
1).
populations
of
time
and
a coyote,
Canis latrans
118
DECEMBER, 1993
zzz
8)
sun
runf s uvd
U
0
cu
06) snIWI
oo
snllvd
0\
I)
I)
vuvnui8.nn snyd/
ap! Q
suaasvrau1a
iumuyavq sn8vpigS
enn Cl
a0
zz
i8apuauvs ruognpnv sn8vju/(
o\
en
O's
ra
v,
pnauuaq
snauroflva
snd7
0
y
8'
K,
r"
0
E
b91)
cv
P
U
00
4.
Lb)
sporavw
sadrasnf vruoloaN
a
0
000
8
ar
1(
6 SO` I)
sadpnu!(
ayaaagsnyydouuads
zU
y
x:
en
y
ea
4.
0
1
U
V
nz' ^ 3^'
zs
CI
00
8v/
C `
U
6
a
a a 414
cI '
i
4"
C'
r r
gg
Z' WC3WrY CI C) .
G/
3 3 o VI
s
a ySo
48
O- '.
Z {_
ri
s =
am.+
C
1 ,.
41 '.:
R.
La Jolla,
clepticus),
rattus,
has
been
not
lagomorphs ( Lewis
survey,
the present
foxi,
although
S.
1988).
et al.
from
reported
During
a. sanctidiegi.
Another
flea, Euhoplopsylla
rabbit
occurs
southern
California being
cinerascens,
arizonae,
Riverside
San
Los Angeles
of this
specimens
flea
were
during
collected
An
et al. 1970).
The capacity for C. f.felis to
transmit plague is probably negligible ( Lewis et al.
1988). Most( 11 of 17) specimens of C.f.fells collected
S.
S.
and
on
( Adams
a.
Most
Rancho Bernardo.
auduboni ( sanctidiegi),
with
from Sylvilagus
records
County,
County ( Augustson 1943).
Diego County record reports it
bachmani
unpublished
glacialis
primarily
survey
the current
two from L. c.
with
bennettii.
In North America, Pulex
be
associations and
P.
example,
of other
a wide
also occurs on
variety
on canids,
and swine(
1985).
Leptopsyllid Fleas
Holland
on mule
deer in
City
of
San Diego,
spp.
La Jolla,
County
Pulex
identified
were
and on
important
plague vector
Lewis
et al.
1988).
In Los
Prince 1943).
were
as a vector
plague- positive
collected
musculus (
Smit
importance
1958),
M.
specimens
as simulans or
and
16 of 28 opossums, mostly
on a coyote,
both host
as to
mammals(
simulans
but
rodents,
Pulex irritans
in the
119
DECEMBER, 1993
among
sylvatic rodents (
specimens
of this
flea
1980). The
from
present
various parts of
plague
the world(
Hopla
of plague (
irritans
The
found
cat
throughout the
Although its
attacks
report
normal
it on 40
of 41 opossums
in the
various
distinct desert
conditions (
Hubbard 1947).
Another
and
September,
120
being
collected
southern
Fifteen
specimens were
1983),
while an
1981
unpublished
and
The
m).
found P. h.
specimens(
P.
of 44)
Nelson
Crestline
San Diego
present
pacifica
41
record reports a
900
DECEMBER, 1993
County
survey
also
Nelson 1983).
occurring in January.
Odontopsylla dentatus occurs from the Rocky
Mountains to the Pacific Coast States( Hubbard 1947).
Its preferred hosts are rabbits and hares and is also found
on their predators,
and
Nelson 1983).
Augustson 1943). An
record reports
with
a single specimen
Diego Zoo.
of
unpublished
it from S. a.
During
0. dentatus
were
San
sanctidiegi, Rancho
occurring
the present
found
on a
on
March ( Schwan
County
Diego County
R.
Utah,
southeastern
1977).
occurs
well as
Diego
California
San
rattus,
to southern
east
and southern
Southern California
Nevada,
Arizona( Barnes
records
et al.
indicate that it
County
records are
from
nests of
N. fuscipes
Barnes
et al.
collected
1977). Most
A.
generally
agree
Lewis
et al.
that A.
1988),
nudatus
current
findings
subspecies
are
infrequently
al.
common on
hosts
during
are
record
1943, Lewis
1988).
et al.
specimens of A. n. nudatus
n. nudatus(
and
Although most
Hystrichopsyllid Fleas
The wood rat flea, Anomiopsyllus n. nudatus, ranges
southern
Nelson 1983).
Bernardo,
Morena.
from
and
121
DECEMBER, 1993
i)
snalwofrlva a spo
Cyy
8
N
6)
snlnasnw
snyy
E
o
i 6) snllve snnvd
tO
suaasveaula
1)
Z)
luvunpvq sn8nilnl'CS
suvinwls slll8v s(
wopodrq
cin
xvllvf fsnglvu8oead
C
Elk)
co
E
a) snpnvat8uol
spop8aui
skwoiuopoegnaJ
r;
1)
lCa7Moa
miCoq snadwoead
Cl
i
yy)
woead
w;
Co
08)
nlaqum8 snvlauvw
snaswwad
CS
snas
Cwoead
o
vi
o
0c
91)
slulsur
o.
Lb)
swaavw
vwo
z
oj
oal l
sadiasnf vwoJoaNN
en
4.
4.
ci.
3i
dU
6"
0
tsR
r(
6S0` i)
sadrpnu!
Kagaaaq
snlrgdouuadg
0.
N. N
to
y
i
F3
1 .. 1,
4.
c-
a .E .
O O p
g.
L.1sod
4.
332 ",
122
Augustson 1943).
County
DECEMBER, 1993
311- 318.
During
examined
in March, 1961,
near
Julian.
the present
cummingi were
Augustson, G. F.
1943.
gentilis, ranges
from
et al.
1988). It is
endemic to the
1988).
It is apparently less
California,
occulatus,
Riverside
common
in
bat
et al.
southern
69- 89.
Barnes, A. M.
Mammals Not
current
of
M.
not
gentilis
Yielding
Fleas
1982.
wish
to thank the
confirmed or identified
following
individuals
who
Entomology, Iowa
State University, Ames, Iowa; Dr. Edwardo G. Campos
of
1989.
efforts and
revisions,
Lastly,
I dedicate this
work to
Shoemake,
southern
951.
ground
squirrels (
Citellus beecheyl) in
Kern
123
DECEMBER, 1993
County,
76.
and
1983.
M. Rothschild.
fleas
ceratophyllid
and notes on
Pp.
Rothschild
tophyllidae:
Cera-
fleas. The
collection of
family
key
relation-
D. S. Longanecker.
andmedical
and
U. K. Distrib.
by
228 pp.
Holdenried, R., F. C. Evans,
1951.
and
mammalian
in
relationships
community in the
of
Canada, Alaska
Memoirs
Greenland ( Siphonaptera).
Entomological
Society
of
Canada -
and
of
the
No. 130.
1949. Fleas
of
1943. Sylvatic
plague
studies.
IV.
1944.
T. Mc Gregor.
Am. J.
and
and
Trop.
transmission
of
J. M. Brennan.
1957.
An
forest
mammals
and
Miles, V. I., A. R. Kinney, and H. E. Stark. 1957. Fleahost relationships of associated Rattus and native
Univ.
and
F. M. Prince.
1956.
Studies
on
124
of
various
studies
species
in California- the
in
of sylvatic
ecology in California.
Conf. 9: 89- 96.
plague
rodents
rodents,
DECEMBER, 1993
in
dissemination
of plague.
700.
1979.
Field
studies.
Part II.
determining seasonal
The ecology
of murine
typhus - a critical
revised
mice, and a
DECEMBER, 1993
ABSTRACT: Vectobac 12AS, a liquid formulation containing the microbial agent Bacillus thuringiensis var.
israelensis, has proven to be an effective larvicide for mosquito abatement. Like other B. t.i. formulations, the
duration of larvicidal activity is relatively short, usually lasting no more than a week. The studies reported here
examined the possibility of extending the period of efficacy for the control of Culex mosquitoes by increasing the
dosage to five and ten times the minimum effective rate. Dosages of 0.2, 1. 0, and 2. 0 kg/ ha of this B. t.i. formulation
were evaluated against natural mixed populations of Culex quinquefasciatus, Culex stigmatosoma, and Culex
tarsalis occurring in field ponds at two locations in southern California. The results showed that the excessive
Vectobac 12AS dosages did not significantly extend the duration of effectiveness for controlling mosquito larvae.
Pertinent to the outcome of these studies were two important natural regulating factors which also affected the
abundance of mosquito populations in the ponds: 1) a progressive reduction in ovipositional attractancy of the
breeding site and 2) an increasing incidence of predation by aquatic macro-invertebrates. As a result of these
interacting factors, the larval populations following treatment with the three dosages of Vectobac 12AS never
recovered, demonstrating that larvicides with a margin of safety to nontarget biota can achieve desirable results using
minimum effective dosages, negating the justification for using higher rates to extend control.
and repeated treatments. In view of the anticipated rise
INTRODUCTION
more
registered
ten
than
years
ago,
being
made
to improve the
continuing
efforts are
B. t. i. in
from,
in
effectiveness of
mosquito control.
With
various commercial
B. t.i. formulations
long- lasting
that assessed
control
various
control of mosquitoes.
In
studies
biweekly)
or
impoundment
and
of
and
commercial
it
was
longevity
B. t.i.
low desert
region),
the major
Darwazeh 1985,
than the
product.
is
much greater
The
purpose of
this study
rates of application,
five
Entomology, University
of
et al. 1982).
and
the
of
Valley (
cost of treatment,
and
Department
Coachella
Mulla
of
determine if higher
the
control of asynchronous
Since the
1990).
and surveillance
cost of material,
efficacy
for the
mosquitoes(
Lacey
monitoring,
was to
weekly
Lacey
126
using
400
ml
located
observed
samples per
to be concentrated.
immatures
Counts
DECEMBER, 1993
were
of sampled
and
4th- instars),
and pupae.
after,
and
minimum of
species
monitored
using
composition,
water
temperature
minimum- maximum
thermometer.
mosquito
in the
rate),
1. 0,
and
2. 0
untreated controls.
used
in
until all
each test.
available, three or
of
et
al.
replicates per
1971)
treatment
31. 3 C.
per
1).
dip ( Fig.
were
calculated
which
compensates
for
in the untreated
natural
ponds.
statistically by using
Larval
indications
identified. Pond
and
basis
post- treatment
weekly
sampling date
was
on
To determine
thereafter.
analysis of variance (
ANOVA).
00.2
a 1.
ponds.
A statistical
kg/ ha
0 kg/ ha
80
2. 0 kg/ ha
Check
60
40
M
ven
O
20
bq
0
0
14
21
28
Days Post-Treatment
Figure 1.
Efficacy of various rates of Vectobac 12AS against Culex larvae at Riverside ponds( AugustSeptember 1991).
between the
comparison
the
significant difference
post- treatment
densities
of
Unlike the
results
variability. There
and
lowest
only three per dip. Not until after day ten ( three days
after the addition of a secondary supplement of organic
material to increase oviposition) did the numbers of
rates,
throughout the
in
Fig. 1) reaching
than
those seen in
higher
that
were
levels
significantly
0. 001 and P
P
<
ANOVA,
rates
(
treatment
other
the
larval densities
days 10
on
had
the high
due to the
was reduced
in the
into these
B. t.i.
dip
per
dip)
per
counts
by
highest
was
following
highest
rates.
This
the high
control,
averaging
progressed,
these and
develop,
to
continued
days 10 and 21 (
resurgence seen at
untreated ponds at Riverside was monitored using 4thinstar larvae to ensure accurate determinations. Initially
( samples taken eight days post- flooding) Culex
quinquefasciatus
early
Fig.
Fig.
2).
those of Cx. tarsalis steadily increased. By day 15 postflooding, Cx. tarsalis had become the dominant species.
Subsequently, on day 22, Cx. quinquefasciatus
1).
Because the
lowest
of
warranted.
the
ponds remained
With few
there was
contributed
were
due to
extended
other
oviposition
factors
activity
consider.
invertebrates
other
little
decline in
evidence
that
present
predators
in the
ponds,
significantly
Fig.
1).
larval
populations
in the
in early
activity was indirectly confirmed by a reduction
instar larvae in the check ponds. Initial samples averaged
dip
leveling
for the
omitted).
entirely
of
with an occasional
of the
and
almost
important to
larvae
larvicidal
densities
population
activity
populations of
highest rate
and the
duration
per
and
kg/ha.
lowest
and
and
producing the
This
days
two
the
As time
dip.
in this
concentration
38 early instars
by
ponds.
complete reduction of
was produced
averaging 4
occurred
middle
indicated
to ineffective levels.
While nearly
instar presence
flow
in the
apparent
treatment.
flowing
flushing and dilution action of
the
water,
additional
rates(
replicated ponds
amounts of water
treatment
already
see
Consequently,
the
21 (
and
0. 01, respectively).
Two of the three
treatment
127
DECEMBER, 1993
Such
off
rest of
a
27 early instars
the assessment period ( data
to an average of
significant
drop
would
make
it
populations to recover to
levels.
observed
samples were averaging only six late instars per dip, not
significantly different from the three B. t.i. treatments.
As time progressed, the densities continued to drop and
on day 14 the experiment was terminated.
128
DECEMBER, 1993
100
Ir
0.
60
40
g.
20
0
8
10
15
22
29
Days Post-Flooding
Figure 2.
Culex tarsalis
Ej Cx. stigmatosoma
Cx. quinquefaciatus
35
Succession of mosquito species( 4th-ins tar) in control ponds dueing first experiment at Riverside
August- September 1991).
100
80
0-
0. 2 kg/ ha
0-
1. 0 kg/ ha
2. 0 kg/ ha
i
60
Check
Y
b
M
40
q+
a
hiiiip,.
g 20
a
0
0
14
Efficacy
of various rates of Vectobac 12AS against Culex larvae at Coachella Valley ponds( October
1991).
129
DECEMBER, 1993
larvicidal activity
There were
was still a
factor.
reasons
several
temperatures were
13. 1
larval densities
first few
primarily
to the onset of inclement weather rather than to a drop in
By day
important regulating
the presence of large numbers of
oviposition
days
of this test.
This reduction
became
factor as
evidenced
hydrophilid
and
by
during
the
was attributed
the
of
ponds.
an
omitted).
Initially,
ponds).
treatment,
6%), but
prevailed
Cx.
two
in the first
species
in the
two
weeks
Cx.
tarsalis
larvae
94%)
and
in November 1991
ponds.
Riverside
quinquefasciatus(
after
trial (
at the
under
Riverside
considerably
facility
cooler
50
0 0.2
o- 1.
kg/ ha
0 kg/ ha
40
9 2.0
kg/ ha
CD
Check
A-
30
M
n
w
20
10 :`
14
21
Figure 4.
Efficacy of various rates of Vectobac 12AS against Culex larvae at Riverside ponds( November
1991).
130
Even
so,
it is apparent
high densities
nearly
zero
levels
in the
as compared to the
during
the
observed
check ponds
declining
the
result
of
populations
decreased
in the
as well as
oviposition
This
activity.
was
confirmed
the
became
and the
low
seen
in Figure 4. Predation
water
of the season
population growth.
Three
Cx.
and
DECEMBER, 1993
Cx.
species,
but because
of the
Cx. tarsalis,
quinquefasciatus,
Cx.
trials,
quinquefasciatus,
which
initially
was
Cx. quinquefasciatus
result of a number of
depends
Cx. tarsalis
to
apparently the
Oviposition behavior
factors.
and
was
the control ponds declined steadily after three days posttreatment, and by day 14 the differences between the
treated and untreated populations had disappeared. All
water
on the
conditions,
quality
organic
especially the
decomposition( Kaul
et al.
presence of
1977, Suleman
and
oviposition
Shinn 1981).
after
This
flooding
Fanara
and
period of
heightened
and
by
strongly attracted to
abundant fermentation but rely on
not
other cues
to
ponds
became
in
our
field
more attractive
the
at
earlier (
October 1991)
this site.
Because
the
at
of
applications( see
day
Fig. 3).
of
predators.
Quickly establishing
duration
of this experiment
ranging from
a mean
131
DECEMBER, 1993
25
20
0-
0. 2 kg/ha
a-
1. 0 kg/ ha
2. 0 kg/ ha
a.
Check
15
A
4
1M
10
Z
00
g
Q
11.'.---
28
21
14
Efficacy of various rates of Vectobac 12AS against Culex larvae at Coachella Valley ponds( March-
Figure 5.
April 1992).
an
Walton
et al.(
The
and
M. S. Mulla.
1974.
Population
1990).
results of
Fanara, D. M.
these studies,
under
the
conditions
advantage
rates(
in conjunction
with
ponds
filled
with
Lacey,
L. A.
and
C. M.
Lacey.
1990.
The medical
nonpolluted water.
and
oviposition
inhabiting
M. S. Mulla.
patterns
in
Temporal
mosquitoes
1993.
Culex
Proc.
eds.),
S6.
132
Mulla, M. S.
and
H. A. Darwazeh. 1985.
formulations
Efficacy
of
DECEMBER, 1993
of
Bacillus thuringiensis H- 14
against mosquito
Environ.
and
its
chironomid midges
DECEMBER, 1993
ABSTRACT: The Oriental latrine fly, Chrysomya megacephala( Fabricius)( Diptera: Calliphoridae), is a blow fly
pest of public health importance that has invaded many new localities over the past two decades. It is a carrier of
numerous food-borne pathogens, including Shigella and Salmonella, as well as infective stages of intestinal
parasites. The distribution of this blow fly has expanded from its original range in the Orient and Australasia to
localities in Africa, South America and, most recently, North America. Its biology, systematics and public health
importance are reviewed for vector control and public health entomologists in the localities that it has recently
invaded. Monitoring the spread of C. megacephala is necessary to minimize its impact on sanitation and health in
the new territories that it invades. Food manufacturing and preparation establishments are at special risk from
increased insanitation attributable to the Oriental latrine fly and must practice good sanitation in order to prevent the
INTRODUCTION
fly
World
throughout the
Within the
fly
1988)
taming
regions
of the earth.
has invaded
and
it
North America,
and
requires
and
its
of the
invading
with
species.
The
blow
fly
is
personal communication).
is known
about this
monitoring its
encourage additional
new environments.
always available
Kitching 1976).
invader( J. C.
review of what
to
published
spreads
unpublished
observations
1U. S. Food&
fly
reports con-
fly
spread
A full
are
a recent
Mariluis,
Peris 1984).
systematics and
the
and
Effective monitoring of an
entomologists and public health
to be familiar
Mariluis
Greenberg
in the Nearctic
range
previously
Desvoidy 1830 (
has
expands
records
fly
scattered
in the
other regions.
professionals
bionomics
warmer
invasion
range
Baumgartner( 1988)
of this species as
Region and
established
coastal counties.
past two
is
expanded
and
collection
Drug Administration, 50 United Nations Plaza Federal Office Bldg., Rm. 526, San Francisco, CA
94102, USA.
2U. S. Food&
Drug Administration, Division of Field Science, Room 12- 39, 5600 Fishers Lane, Rockville, MD
20857, USA.
Grove, CA 92643, USA.
3Orange County Vector Control District, 13001 Garden Grove Blvd., Garden
134
windows
Anterior
cornua wide.
processes.
on
2- 8,
on
9,
spiracle with
spinal
dorsal
10- 13 fingerlike
complete
ventrally; bands on 2- 8
Posterior spinal bands (
dorsally
Spines
than
on
and ventral
Anterior
scar.
usually present
between dorsal
on
6- 10,
6- 11)
band,
anterior pleural
incomplete
dorsally
Papilla P2
and
Holloway
closer to
P1 than to P3.
Body
lacks
fleshy
1- 11.
The genus is
11 but faint
ventrally.
but fewer laterally
complete on
lateral
with
segments
DECEMBER, 1993
An
United States.
cephalopharyngeal skeleton.
The
puparium
spiracles and
which
are
by
may be
recognized
by
the posterior
retained
stage (
Kitching
1976).
The
adult
is
blow
a robust
fly,
8 to 12
body
mm;
Abdominal
Lower( thoracic)
DISTRIBUTION
Male
long
portion and a
1.
2.
setulae.
facets larger
distinct line
of
than
facets
demarcation
Prothoracic
spiracle white
Prothoracic
spiracle
on
lower
across
the
dark
Legs
metallic black;
parafacials( buccae) yellow to orange, at least anteriorly; wings hyaline
in both
front femur
sexes;
hairs
Legs testaceous yellow in part; parafacials orange, densely yellow dusted; Front femur of
male with
3.
4.
dense
erect white
Prespiracular
seta present
Prespiracular
seta absent
hairs
dorsally
Body stout, but rather elongate, submetallic dark blue; thoracic squama largely blackish
Chrysomya pacifica Kurahashi
Body stout and rounded, metallic greenish blue; thoracic squama largely brownish
Chrysomya
megacephala(
Fabricius)
DECEMBER, 1993
based on the
finding of a"
form there
natural"
and
in the
was a separate
Kurahashi 1991)
is
from C.
species
natural"
C. megacephala
Mexico(
James 1971),
Greenberg
Park
peninsula(
Palearctic in Afghanistan
Kano 1963).
and
It is found
Indian Ocean
on
Hawaii(
1940)
and
Malaysian
far
as
east
earlier (
1960)
inclusions
range except
specimen
adult male
flies, and Cole ( 1969) reported that the genus did not
1988).
Oriental latrine
by
fly
Shewell' s ( 1987)
also
Baumgartner
and
Greenberg
1984).
A single male
Thecontemporary patternofdispersal
of this
first
an area
a new
territory
consists of
1978, C.
North America.
to other
modern expansions of
continent
western
the
in
dispersal
C. megacephala into
where
occur
contributes to the
Greenberg
1961
1984).
Greenberg
of the
trade (
and
spread
1955).
Prior to
range.
outside this
known from
was not
Baumgartner
The
for
in the
transportation
Mexico.
also
1977, C. megacephala
will provide
as
Korea
and
Hardy
Greenberg 1988,
islands in the
and on the
of
megacephala,
yet unknown.
and
135
was
found
on
spp. have been collected in the New World since the last
the African
century ( Baumgartner
was
where
found in
it is now
Greenberg
1984),
those
South
in inland
coastal
widespread
and
it was found in
same year
the
of
Brazil
occurred earlier (
do Prado
and
GuimarAes
from Angola
during
an exodus of
et al.
1979, Peris
1987),
Paraguay(
Barrios
Ecuador( Olsen,
and
by
1981b).
et al.
fly
was said
subsequent
blow
to occur in
fly
surveys
being
136
DECEMBER, 1993
ORIENTAL
OCEANIA
AUSTRALIAN
China
Australia
Carolines
Afghanistan
Angola?
India
N. Zealand
Fiji
Iran?
Canary
Indonesia
Papua/ N.G.
Gilberts
Japan
Egypt?
Ecuador
Malaysia
Guam
Korea
Ghana
Paraguay
Philippines
Hawaii
Malagasy
Peru?
Sri Lanka
Marianas
Mauritius
Venezuela
Taiwan
Marshalls
Senegal
Thailand
Palau
Seychelles
Viet Nam
Samoa
South Africa
I.
Argentina
Mexico
Brazil
U.S. A.
Solomons
Tonga
Vanuatu
indicates that
latrine
few
to monitor
used
fly
was
collected
1990),
in
and
urban
C.
megacephala
Southern California
has
since
females
and
been
truck from the Los Angeles area. It is likely that the San
Diego flies were transported south from Los Angeles in
were
Sidebottom
in
collected
Altadena,
of Sciences.
the
communities
Long
of
Beach, Orange,
bring
California
three coastal
Angeles, Orange,
counties (
Orange
The
immatures
counties
year
likely
have
examination
dumpsters)
Ysidro
and
of
20
was made
Otay
in the U.S. is
have
fly
occurred
in August,
sites ( meat
colonization
originating from
western
Mexico.
An
market
in San
border crossing
of San Diego. No adult or larval
Mesa, the
may have
Mexico, an
megacephala
attractive
in
An overland
or shortly before.
To determine whether C.
across
uncertain.
1988
invaded California
are
to
authors
in California
and adults
every
arrived
and
Los
principal
is
DECEMBER, 1993
Pacific invasion
mutes that
include
megacephala
have been
Shanghai
Burbank is a
Other
for C.
Honolulu
route
to
pan-
postulated
personal
Kong to California
communication), both aided by
Espin
Pavan 1983).
These
and
Schnack 1985)
or
the chief
and
reports
postulate of a
Pacific invasion
There is
a need
objection
modern
C.
137
not occur
support
for California.
route
now,
megacephala
decades
the
is why the
ago.
Another
pathogen Staphylococcus muscae may cause early dieoff of males as soon as 15 days after emergence( Mariluis
of
C. megacephala is described by
cycle
and
from egg to
adult is
or
consists
of
two to six
nonfeeding
to pupate( Bohart
order
and
Mature larvae
from the
feeding
Gressitt 1951).
enter a
site
The
in
Mohanan
non-
1980b).
larval
prepupal
prolonged
numbers of
while
on
ratio
days to
complete
larval
to
density and
density. Large
periods of
over seven
and
larvae
in inverse
in larval developmental
result
less,
is partly dependent
stages
may be
stage
five days
conditions
food
or
may take
development( Goodbrod
appears
to be
body
along
food
larvae that do
source
Greenberg
The
is
exhausted
do
before
1991).
pupal stage
longevity
for both
days,( Subramanian
to
20
and
In
adults
laboratory
is nearly 1: 1( Subramanian
and
Mohanan 1980b,
and cadavers(
Goff
et al.
with
and
strategy
larvae from a
Das
1981)
a survival
and
larvae become
facultative
agents of myiasis
Mariluis
and
Schnack
1985),
exophilous
138
1971)
Greenberg
and
synanthropicinbehavior.
fly,
it is
1951,
attracted
Roy
and
1990). It is
also attracted to
excrement ( Jettmar
both indoors
summarizes
been
and
where
Gressitt
Sidebottom
( Schnack
it breeds in
et all.
and to corpses
Goff 1991).
settlements
and
positive
values
reflect
progressive
and outdoors (
The degree
different
latrines
and
TABLE 2
reported
in human
to hemi-
eusynanthropic
DECEMBER, 1993
of
synanthropy,
settlements, varies
geographical
areas.
or success at
for C.
megacephala.
megacephala
measure
surviving
in
has been
Similar
ATTRACTION
Foodstuffs
general
eggs
U.S. A.
fish
Brazil, India, Malaysia, Palau, S. Africa, Sri Lanka, U.S. A., Venezuela
fruit
meat
melons
China
shellfish
sweets
Structures
houses
markets
latrines/ toilets
restaurants
Mexico, U.S. A.
ships
slaughterhouses
Sri Lanka
Other
carrion
Australia, Brazil, Guam, Hawaii, Malaysia, Papua/ New Guinea, S. Africa, Sri Lanka
corpses
feces
garbage cans
landfills
Malaysia,
Paraguay, U. S. A.
Mariluis et
less
with
of
al.
mathematical observations of
C. megacephala for
are consistent
the preference
habitats from
( Bohart
1991) to
apartment
complexes
in Hawaii
Gressitt 1951).
( Bennett 1989).
Wilton 1961).
we
and
Brazil( do Prado&
Holloway
139
DECEMBER, 1993
larval activity
during
pupal stage
Chrysomya
species,
in
puparia
temperatures as
of cooler
C.
areas,
Das
is
prolonged
is the
In
with
This may
during periods
Wiedemann), in
central
bimodal
per
exhibit
two
population peaks
and Dasgupta
response
populations
megacephala
fluctuations
seasonal
protected sites.
albiceps(
be in
year(
of adult and
FOOD SANITATION
1975).
There
are
natural
of
a number
enemies
of the
larvae( Tullis
McCook
was
pupae near a
and
California.
predaceous
histerid beetles
flesh
larvae ( Diptera:
carrying
dumpster in Orange
observed
Coleoptera: Histeridae)
and
off
live larvae
County,
fly
cited
previously
life
Wells
al.
1981).
We
and other
observed predation
both
species are
Pacific island
spider,
Vespula
of
Nephila
by
V.
ones.
fly
longevity ( Ho 1990). Vertebrate
phoretic
on
feed opportunistically
1981),
On the
Fabricius) ( Araneae:
on adults(
Public health
Orange Counties,
reduce
Down 1946).
Oriental latrine
and
et
maculata (
Harris
germanica on
recently introduced
Palau,
megacephala(
preys
and
Hymenoptera:
yellowjacket,
C. megacephala
on adult
1992). In the
Greenberg
not observed
germanica(
Chow 1940).
and
history
may
insectivores
Gressitt 1954).
and
disease ( Kurahashi
and
Banu 1989),
and
140
Because
handling
the
area
be transported from
will
very dangerous to
places
pathogens
they may
places where
not
be
where
megacephala
Bohan
and
is considerable for
a number of reasons
by
when attracted
various
food
buildings
enters
products and
is
also
DECEMBER, 1993
is
not
demonstrated.
This is because
Bandler 1990).
extraction
items that
unsavory
environments where it
it easily becomes
transmitting its full
As
thrives.
in
mired
occur
in the
a rather
food
urban
clumsy
animal,
dies there,
and
process.
its
to
potential
for
transmitting
pathogens
from
megacephala
from facilities in
of
food
illustrates
identified
to
The
identify
these
fragments ( Kano
and
Sato 1951).
The
reservoirs
C.
the
exclude
order to prevent
CONCLUSION
and
Ueta ( 1990)
megacephala to reduce
from
and
et
control of
al.
C.
by
industry
fly. Studies of
the food industry
the
Sulaiman
for the
certain pesticides,
1988).
it is necessary
to
measures to protect
rely
on prevention
foods from
and exclusion
contamination
by these
control(
foods
long- term
solution to
Esser 1991b).
fragments
were
identified from
by local
authorities,
among them
being
the presence of
baked
to invasion
by
this filth
fly.
Adequate
goods.
Over 25 live
adult
flies, mostly
Observations
such as
DECEMBER, 1993
Nicholas J. Richter
and
J. Richard Gorham(
and
141
FDA),
R. L. Berstein, San
all
Francisco State
in the tropics.
We
also thank
Juan C.
of
Medical
National Institute
Entomology,
University
and the
1988.
Spread of introduced
of
Wheeling,
Baumgartner, D. L.
of
Operations
B. Monzon, Mahidol
University ( Thailand);
Demei
and
who
in California),
Woodley
first discovered C.
and
the many
other
H. G. Davis.
1981.
The
yellowjackets of
Isolation.
Calliphoridae( Diptera).
of rodent- baited
Avancini, R. M. P.
breeding
responsible
and
Manure
and
Coleoptera)
caged
layer hens. J.
M. T. Ueta.
insects ( Diptera
for
cestoidosis
in
1990.
312.
California Department
of
Food&
Agriculture.
1988.
Chow, C. Y.
1940.
142
bacteria in
Peiping,
153.
DECEMBER, 1993
Berkeley.
360 pp.
Ferreira, M. J. M.
1978.
Sinantropia de dipteros
of
the North
blow- flies
and
and
B. Dasgupta.
1982b.
Sex- ratio
of
1991.
133.
Dear, J. P.
Dear, J. P.
Fauna
of
DSIR Publ.,
Dowell, R. V.
and
in
1986. Arthropod
succession
O' ahu,
island
of
531.
1990.
influencing
oviposition,
mortality in Chrysomya
Diptera: Calliphoridae), a pest of
growth
megacephala (
salted
Factors
and
dried fish in
south- east
1989.
fish
industry
in
south-
measures reduce
Gorham, J. R.
1991.
NY.
blowfly
of two
closelyChrysomya
Greenberg, B.
1973. Flies
and
disease. Vol. 2.
Biology
and
143
DECEMBER, 1993
Holloway,
B. A.
1991.
Identification of third-instar
Greenberg,
1988.
B.
Chrysomya
F.)
megacephala (
and notes on
species present
in the New
Illingworth, J.
F.
megacephala (
1926.
Notes on Chrysomyia
Fabr.) ( Diptera).
Proc. Haw.
Greenberg,
1991.
B.
Greenberg,
B.
and
1984.
M. L. Szyska.
Immature
of Micronesia
1: 165- 166.
Chrysomya Robineau-
74- 85.
1979.
Dispersal
introduced
and
species
distribution
of
of three
of
Gunatilake, K.
and
M. L. Goff.
1989.
Detection
organophosphate
5: 465- 474.
Hall, D. G.
1948.
The blowflies
of
Kano, R.
and
K. Sato.
1951.
Hardy,
of
146.
Kitching, R. L. 1976. The immature stages of the OldWorld screw- worm fly, Chrysomya bezziana
Villeneuve, with comparative notes on other
Harris, A. H.
dissemination
parasites
by
1946. Studies
H. A. Down.
and
flies in
various
of
the
human intestinal
localities
on
Guam.
Australian
species
Calliphoridae).
of
Chrysomya ( Diptera,
66:
195-
203.
1990.
peninsular
between
Macrochelidae)
61- 68.
association
Atari:
Macrocheles
in
Phoretic
muscaedomesticae
and
144
Fabricius) newly
recorded
from
DECEMBER, 1993
Singapore,
with
Chrysomya from
species of
notes
on
C. defixa ( Diptera:
1981.
descriptions
of
with
fly Chrysomya
megacephala,
of a synanthropic
Spinelli.
in New Guinea
1990.
698.
Meng, H.,
Kurahashi, H.
1991.
with
N. Kuo, P. Wei,
and
C. Kao.
megacephala (
Kurahashi, H.
and
Q.
Banu.
1989.
Notes
on
the
1966.
Mihara, M.
and
Fab.)
and
Ch.
H. Kurahashi.
phaonis (
1991.
Seg.). (
in
Base- line
megacephala (
Microanalytical
expansion of the
range of
Trop.
of
Samoa
para La
Argentina, Bolivia y Ecuador( Diptera). Rev. Soc.
1- 49.
Oldroyd, H.
1964.
Weidenfield&
and
para una
observations on
Chrysomya
megacephala(
Fabr.)
145
DECEMBER, 1993
and the
Wied.)
130.
and
T. H.
85- 91.
Chrysomya
megacephala (
280- 281.
1960. Studies
Park, S. H.
of
374.
Kyung- Pook
Park, S. H.
on
and
collected
Patton, W. S.
and
E. C.
and
J. Smart.
1940.
Penis, S. V.
Chrysomya
en
world genera
McAlpine,
ed.).
Soc.
Esp.
Bol. R.
and
R. Kano.
1979.
and
J. H. Guimaraes.
de dispersAo
1982. Estado
distribuicao do
genero
Spradbery, J.
P.
1979.
Discovery
of the
six
South
Calliphoridae)
Ann. S.
and their
immature
stages.
and
A. R. Sohadi.
146
1988a.
Isolations
DECEMBER, 1993
bacteria
of entempathogenic
including
the
taxonomic
keys ( Diptera:
and
R. Iberahim.
243- 272.
Entomol. 2: 1- 6.
and
parasite
J.
Jeffery.
burdens
on
1989.
Cyclorr-
sundrying.
625- 629.
and
permethrin
against
filth flies
Wells, J. D.
and
B.
Greenberg.
1992.
Interaction
at a garbage
Lower
reaches of the
1951.
1957a.
Chrysomyia
Hemipyrellia Tnsd.,
southeastern
Wijesundara, D. P.
R- D.,
and
Lucilia
R- D.,
Calliphora R- D. from
Wijesundara, D. P.
carrion
in
succession
a tropical rainforest on
O' ahu
and
R. Kano.
1979.
DECEMBER, 1993
ABSTRACT: Northeastern Greece contains four of the eleven large wetlands of the country that are protected by
the Ramsar Convention. Malaria was once a serious health problem, but intensive antimalarial measures eradicated
the disease in 1965. DDT residues are still traceable in many components of the food chain. The destruction caused
in the sensitive ecosystems by the chemicals and plans for tourist promotion of landscapes prompted us to adopt
control measures that would be less harmful to the environment. A pilot study using biological control with Bacillus
thuringiensis var. israelensis( Bti) began in the Summer of 1992. Results revealed 95- 100 percent mortality in the
2nd and 3rd larval instars 24 hours following the application of Bti in concentrations of 250 gr/ha. In a parallel study
on mosquito species, 27 species in five genera were found in the area.
INTRODUCTION
The
coastal
by
characterized
about
region
of
Northeast
Greece,
rivers,
45
Km2),
and
irrigation
agricultural
system
Evros Delta
to the wetlands
in the
constructed
development
by
prohibited
15 Km2),
and numerous
is the
of
the tremendous
World War.
in the
health
temporary breeding
permanent or
wetlands.
Malaria
problem
in this
human
especially DDT
for
carbons and
were
the
malaria.
prohibited,
in mother'
et al.
organophosphates,
first intensive
vector
and
control
use of DDT
measures
indices( Ouzounis
detected
Test of Bti
insecticides
used
using
WHO 1986).
In this preliminary
work,
the
possibility
of
of
148
TABLE 1.
DECEMBER, 1993
Habitats Created
Physical Characteristics
m2
surface size/
depth/ cm
II
III
20000
200
100
500
5000
40- 60
30
50
10- 15
10- 20
larvae/ 1 H2O
pH
air temperature
D. O. mg/ 1
D. O. percent
mS/ cm
conductivity
salinity/
150
20
50
10- 15
10- 20
8. 30
8. 32
7. 62
8. 81
8. 49
30
30
18. 5
19. 1
32
27. 5
25
17. 4
23. 8
30.2
10. 5
4. 9
1. 8
10. 6
9. 1
133
62
29
130
120
5. 65
5. 09
0. 75
8. 61
0.32
3. 5
3. 1
0. 5
5. 4
0. 1
temperature
water
IV
00
first
preparation
habitat IV),
in the
second one
abundant.
was applied
The
July- August
habitats
other
was a
1991
and
during
May-
The larvae
their natural
used
in the tests
breeding
There
application.
lacking
which were
were collected
places prior
from
to Bti treatment,
by
instar
cages at the
five cages,
spaced
larval habitat,
depending
Containers of equal
filled
with
in TABLE
an equal number of
Larval
1.
density
and
V,
and
of
Bti
these habitats is
Anopheline larvae
used
of
the
Aedes larvae
of
prior to
same
Study
habitat
containing
to serve as control.
in habitats I
treatment and
given
volume, with
from the
were used
in the
habitats.
Mosquito Species
Sixteen
Bti Efficacy
Larval mortality was recorded 24 hours after
larvae,
water
were placed
on
Mosquito Species
for the collection
of
DECEMBER, 1993
in the
July
and
August, the
collections was
in TABLES 2, 3,
and
Anopheles
maculipennis
4.
During
in daytime
followed
by
coastal
were
TABLE 2:
149
Species
Relative abundance*
Anopheles
maculipennis
Anopheles
sacharovi
Anopheles
superpictus
Meigen
abundant
Favre
abundant
Grassi
moderate
rare
Anopheles
claviger
rare
Anopheles
algeriensis
abundant:>
Meigen
Theobald
rare
TABLE 3.
Mosquito species
Aedes caspius*
Aedes cantans( Meigen)
Aedes vexans( Meigen)
Aedes geniculatus( Olivier)
Aedes annulipes( Meigen)
Aedes pulchritarsis( Rondani)
rare;<
25
mosquitoes
in
average
in 15
min.
150
TABLE 4.
DECEMBER, 1993
Relative abundance
species
Aedes
caspius(
Pallas)
Aedes
vexans(
Meigen)
Aedes
cantans(
Aedes
geniculatus(
Aedes
echinus(
abundant*
moderate**
Meigen)
rare***
Olivier)
moderate**
Edwards)
rare***
Anopheles
abundant*
Ficabli)
modestus(
abundant*
Meigen)
maculipennis(
rare***
50 mosquitoes in 15 min.
in the
July( TABLES
middle of
Human baits
were
used
for
and
outdoor
3).
evening
5.
Acknowledgments
human
biting
very
species
modestus and
frequently collected
in the
coastal areas.
Aedes
in the
vexans were
study.
appreciated.
CONCLUSIONS
REFERENCES CITED
1.
for
use
in Greece.
Stechmuckenbekampfung. BiologieinUnsererZeit
4: 10- 111.
2.
densities
maximum
in the area
and attained
during July
and
August.
Betzios, V.
1989.
3.
area.
Anopheles
sacharovi
followed
An.
in
winter.
4.
in
frequently
Margalit, J.
and
of
Bacillus
DECEMBER, 1993
thuringiensis var.
israelensis. J. Am.
Mosq. Cont.
151
1578.
Assoc. 1: 1- 7.
Pantazis, G. 1935. La faune des Culicides de Greece.
Ouzounis, K.
1990.
Biocides in
mother' s
milk.
of
and
M. Bahadir.
368.
ausgewaehlten
chenlands (
Umweltproben Nordgire-
DECEMBER, 1993
ABSTRACT: We examined the distribution of adult resting mosquitoes at 30 sites surrounding a residence in the
urban Los Angeles area. The distribution of adult resting mosquitoes remained remarkably constant throughout the
five month( June- October) study; resting adult mosquitoes consistently preferred sites on vertical surfaces above
decaying vegetation. The number of mosquitoes at resting sites was greatest at midday( 1100- 1300 PST) and was
less at sampling periods in the morning( 0700-0900 PST) and late afternoon( 1400- 1600 PST). The abundance of
resting adult mosquitoes at this residence was greatest in June and was lowest in August. Resting mosquitoes were
predominantly males( 61% of the population) and the preponderance of males in AFS Sweeper samples may reflect
the proximity of a larval developmental site to the residence. Biotic factors, such as predators, non- culicine dipterans
chironomids), or other mosquito species did not explain the observed distribution of the dominant mosquito, Culex
quinquefasciatus, and the distributions of three less abundant species. Differences among sampling sites for abiotic
factors, such as light intensity, ambient air temperature, and relative humidity did not satisfactorily explain the
observed distributions of resting mosquitoes; however, the importance of within- site differences ( microscale
differences) in these factors may have been underestimated.
INTRODUCTION
Studies
mosquitoes
of
in
the distribution
nature are
of
resting adult
limited. Service( 1969, 1971)
of
hosts
influenced
by
Previous
shade.
Bidlingmayer( 1969)
and
studies
by
Service ( 1969)
Edman
and
showed that
blood- fed
al. 1963).
arachnid predators
deterred Culex
negative
binomial
model.
mosquitoes.
intensity
As
compared to
was of greater
importance in the
selection of
Schreiber
et al.
1989).
2Current address: Florida A&M University, John A. Mulrennan Sr. Research Laboratory, 4000 Frankford Ave.,
Panama City, FL 32405, U.S. A.
3Current address: College ofLife Sciences and the Department ofZoology, University ofMaryland, College Park,
MD 20742, U.S. A.
distribution
spatial
and species
to examine
and quadratic trends were also run for the time of day
mosquitoes
landscape
via
to elucidate the
residence,
urban
at one
undertaken
distribution,
management strategies
by
and
architecture
matrices.
adult
reducing
harborage.
mosquito
153
DECEMBER, 1993
Adult resting mosquitoes were collected semimonthly from June to October, 1989, at 30 sites
in Irvine, California.
a residence
surrounding
The
predators ( spiders
detailed description
of the
included trees,
i. e.,
sites
structures
The 30 sampling
above
and
walls,
flowers,
2)
sites
3) flowers(
eaves, walls);(
backyard
vegetation (
above unmulched
beds);( 4)
Fig.
from
were taken
ground level to
m and
Fig.
vegetation(
Twice
5) front
Vertical
1).
m above
vegetation.
an
ants),
its
site and
study
and
insects.
residence was
a more
nondipteran)
Samples
from 4
m to
in the backyard
Least square
GLM
procedure,
1).
month, all of the
during each
resting sites at
three time intervals:
Twenty-
and
chosen
for
each
period,
sampling
with
samplings
sequential
eliminating
sampled was
thus,
temporal
bias.
A
repeated measure
ANOVA
RESULTS
for
Because
among resting
to
in
beds
on most
analysis.
analysis.
day)
in the flowers
The
Habitat
and the
within- subject
was
catch(
TABLE 1).
number of
soma(
13%),
habitats).
categories (
above unmulched
prior
site
was
log+
examined
value
as
interaction terms
were
tested as
and
Culex
tarsalis(
months).
TABLE 2).
154
DECEMBER, 1993
D)
14;.; .
0
0
'
FRONT
CC ;
ofSe. . ....
8>
1:3
C
o
0 CS
CD :
z as
rij a
crs 45
E o
tri co . >
a)
E ....
It15i:'
N-1
0)
u)
o
Mitgion
I
N
N
0
0
0
0
1>
as
ON:
3
gtirg!'
IA.
Lk.
v.......-
v...;
laiiii0'"
v.
i
c...
is..
c
c.;:
7*
0
0.
1
Figure 1.
40L" .Cn
i: k.
BAC K
Location ofstratified
sample samples
sites surrounding a residence in Irvine, California( A-Z and FF, QQ, ZZ, WW).
Vertically
were taken
from 0 to 1
m and
4 to 6
m at sites
D, E,
and
F.
155
DECEMBER, 1993
TABLE 1.
Percentages of resting mosquito species and males collected at three discrete time intervals
at the Irvine, California site in 1989.
Species
Culex
Culex
Culex
Culiseta
quinquefasciatus
tarsalis
stigmatosoma
incidens
Time
N1
period
Males
0700-0900
131
63. 4
57. 3
18. 3
16. 0
8. 4
1100- 1300
168
61. 3
64. 9
17. 9
12. 5
4.8
1400- 1600
104
58. 7
65. 4
19. 2
8. 7
6. 7
Total
403
61. 3
62. 5
18. 4
12. 7
6.4
Percentages of resting mosquito species and males collected monthly at the Irvine, California
TABLE 2.
site in 1989.
Species
Culex
Culex
Culex
Culiseta
Males
quinquefasciatus
tarsalis
stigmatosoma
incidens
June
64. 8
40. 7
27. 1
18. 6
13. 6
July
84. 2
65. 6
14.4
8. 9
11. 1
August
61. 7
74. 2
17. 7
8. 1
0. 0
September
58. 1
65. 1
14. 3
19.0
October
51. 9
78. 6
17. 9
3. 6
Time
period
The
percentage of males
was, on average,
of males
in
in AFS Sweeper
samples
samples
did
not change
However,
appreciably across
between 62
1. 6
0
in
in
84
percent and
percent of the
The
males were
individuals
proportion
collected
of males
4-6
m:
0- 1
is =
Males
were collected
and
mosquitoes at
ts=
resting
among
0. 024).
in
the slope
2. 541,
sites
ts:
P> 0.05).
number of
aspirator samples
months(
P < 0. 01),
until
sex
upper stratum(
diel
more
were evident at
stratum
were
abundant than
Seasonal
in the higher
differed significantly
F4, 14= 3. 93, P=
significant quadratic
3).
sites
156
DECEMBER, 1993
June
July
5
Aug
8
Sept
a\
Oct
_
1
Time of Day
Figure 2.
The average number of mosquitoes collected in AFS Sweeper samples from all habitats from June until
October 1989.
of
morning
and
late
afternoon
site A).
had to be
pooled
Particular resting
in
most
instances.
by resting
DISCUSSION
individuals
collected
significantly
during the
mosquitoes at
was
resting
five
sites
months,
the distribution of
the residence
surrounding
Fig. 3).
1)
a wall above a
bed(
site
FF);
leaf mulch
contained a
layer
of
4)
and(
surface
DECEMBER, 1993
157
Month
Time
June
0700- 0900
10. 52**
1100- 1300
81. 03***
1400- 1600
6. 02*
July
August
September
October
d. f.t
period
2$
0700- 0900
1100- 1300
1400- 1600
0700- 0900
1. 32 NS
1100- 1300
3. 69 NS
1400- 1600
1. 40 NS
0700- 0900
5. 99 NS
1100- 1300
6. 86*
1400- 1600
0. 33 NS
0700-0900
1100- 1300
1400- 1600
3. 93 NS
10. 94**
3. 75 NS
1. 07 NS
13. 17**
0. 48 NS
td.f.were first pooled by number of species so that not more than one of the expected
frequencies was less than five.
Goodness
of
fit test
where
NS
0. 005 <
0. 001 <
P 50.005.
P> 0.05;*=
0. 01< P 50.05,
distributed.
vegetation
metropolitan area(
Schreiber
et al.
than
did the
vegetation
Adult
in front
of the
house.
mosquitoes
crepuscular
periods.
We
sampled
next.
adult
resting sites are not limiting, and these species are not
competing for resting sites around this urban residence.
unambiguously
resting
among
months
in June and
greatest numbers of
from resting
sites
catches suggest
sites
collected at the
Irvine
The
developmental
site was
in
close
proximity to
DECEMBER, 1993
158
1-
O
o
a)
1. 4
v o
o)
13
to
to
a)
>,
CD
0) ao
a)
a)
f0
E C
Z.
E
as
y l..
CD
ti
Qtola
CIEBEND
I
l
0
Potemo
I
0
Lt)
0
V
u7
LO
cn
a)
a)
LL.
O)
gg
1
D)
O_
el
t -
Mr
f-
z
J
CO
C7
r._Y
r m
yiff::::rS:i:; S:::=:::
s: b:>:.:
Fir:2. 2:: ir :
O -
- tA
0I
U)
r
O
V
It)
Co -
ig5n1?j.
ioqumN a5B.)aAV
Figure 3.
Abundance of adult resting mosquitoes collected from June until October 1989 at sampling sites
surrounding a residence in Irvine, California. The numbers of individuals collected at each site were
summed across time periods
in
for
Schreiber
the residence.
percent of
159
DECEMBER, 1993
et al. (
the individuals
1989) found
85
that
breeding/ developmental
freeway drains were males.
near mosquito
sites, such as
resting
adult mosquitoes, a
factor at a particular
of
at a central
location
maybe sufficient to
sampling
sites,
but future
factors
within, and
sites.
among, resting
abiotic
Gardening by
Martinez.
adult mosquitoes.
246- 250.
of sites
vicinity
and
diel
and
pivotal roles
in resting
site
To
understand
have
urban settings, we
mosquito
selection.
distribution,
several residences (
detailed
emphasized
abundance, and
Schreiber
studies of
biting
habits
at
et al.
and
J. E.
Hazelrigg.
1988.
residences
determined.
population
at
resting
remarkably
at individual
the
Irvine
residence
was
better understanding
of
less
predictable.
S. Mulla.
modeling
disease transmission
and
populations
and management of
pestiferous
in the neighborhoods
and,
mosquito
eventually, in the
Service, M. W.
1971.
Acknowledgments
8: 271- 278.
assistance
in the field.
Procedures
REFERENCES CITED
Edman, S. D.
and
of
W. L. Bidlingmayer.
blood- engorged
capacity
News 29: 386-392.
of
Statistics.
Mosq.
mosquitoes.
127.
DECEMBER, 1993
ABSTRACT: A fizzy tablet consisting of a spray-dried powder ofBacillus thuringiensis subsp. israelensis( Bti) was
constructed to obtain a product for mosquito larval control that was easy to store, carry, and use. The formulation
was tested in the early spring of 1991 in 10 small, temporary spring ponds containing mosquito populations ofAedes
cataphylla and Aedes cantans. Half the tablet dose per 5 m2 killed 85 to 88 percent of 2nd to 3rd-instars. A full tablet
dose per 5 m2 killed 93 to 100 percent of 2nd to 3rd and early 4th-instar larvae in ponds without dense vegetation,
and 75 to 82 percent of the same instars in ponds with dense vegetation. In ponds tested later in the season and
consisting nearly of only 4th-instar larvae, the effect was 73 to 79 percent. Dense vegetation or a large amount of
dead leaves on the surface prevented the particles from spreading. The tablets are thus most usable in breeding sites
without these obstacles, such as water reservoirs, garden pools, swimming pools, and small streams. B ti does not kill
late 4th-instar larvae because they do not ingest it.
INTRODUCTION
black
for the
used
larvae for
fly
Sweeney
and
control
of
Product Formulation
10
years
1983).
The
more than
produced
The
production of
B ti is based
process,
a concentration
followed
by
either
formulation
of the
spray- drying
and
and
crystals
formulation
fermentation
on a
spores
The
or a
products
flowable concentrates
formulated by the
be
applied
by helicopter
over
Mosquito larvae
ponds and in
et
al.
1991).
To treat these
small water
are not
very
convenient.
the
sites,
traditional
A better product
for
application,
described for
a similar
Preliminary
lations in
field trials
be easily
tablet ( Becker et
18 to 160
m2)
fizzy
1991).
al.
in
IPS82 lyophilized
Bti formulated
be
Laboratory experiments
using buckets
used in water
containers.
breed in very
formulations
ideally
often
Bioassay of Toxicity
Ponds
formu-
forest.
applying
Novo Nordisk Plant Protection Division, Product Development, 2880 Bagsvaerd, Denmark
between 6 C in
12 C in
mornings to
161
DECEMBER, 1993
some afternoons
during
some
April,
1991.
Sampling
Samples
10 to 20
dipper 30
according to
pond size.
cm and
ponds,
after
to
returned
the ponds after counting, but one to two samples from all
the effect
was
75 to 82
percent (
TABLE 2).
In two
ponds
were brought to
the
period.
Treatment
Ten
ponds were
tablets per
or
m2.
dosed
For their
in
use
ponds with
water surface,
dense
dividing
The
grass
two
was used
drawing
scale
by
of
RESULTS
Toxicity
The
the primary
laboratory bioassay
of ca
potency
production
activity
of
of
powder.
Mosquito Species
Aedes
ponds
and
found.
in
cataphylla and
only
In the
few
Ae.
cantans
other mosquito
beginning
of
all
the guidelines used for Skeetal FC( 600 ITU/mg, 21/ ha).
were
were
dominated in
species
mostly
Monoclonyx sp., a
found in two ponds.
of
The
several ponds.
Field Effects
results obtained
in the
and
ponds
depended
on
162
TABLE 1.
DECEMBER, 1993
Temporary ponds, circular with depth of 10 to 30 cm, area ca 115 m2, were treated with 25 Bti tablets
or ca 1 tablet per 4.5 m2, which is approximately half dosage. The overall result is a reduction of the
larvae population by 83 percent. The raw data are shown to illustrate the inter-sample variation.
Instar
Pretreatment
Sample No.
2.
3.
Post- Treatment
4.
All
3.
0
1
11
10
13
Total
44
59
0. 7
10
Avg./ Sample
6
1
12
3. 7
All
0
0
1
4.
5
6
0.6
1
11
4. 9
Reduction (%)
TABLE 2.
2.
0. 1
0. 5
0. 2
88
85
0. 8
60
83
The table shows the number of larvae caught in three larval stages and the total per pond. The first four
ponds had a water surface without grass or other obstacles and the next three had a water surface full
of grass or dead leaves. These seven lakes were treated early in April when there were very few 4thinstar larvae. The latter two ponds were treated late in April and were dominated by 4th-instar larvae
and pupae( not shown).
Late 4th-instar larvae and pupae do not eat and are therefore not influenced
Pond
Pretreatment
Post- Treatment
2.
3.
4.
All
Reduction
2.
3.
4.
All
(%)
Clear
10
28
168
23
219
13
14
94
Water
10
22
21
47
94
Surface
10
44
25
76
93
Early
20
27
100
Grass
10
44
21
74
and
12
78
77
14
169
15
Leaves
10
26
20
33
79
47
48
10
44
44
13
82
23a
40
76
12
20
75
10
10
79
12
12
73
Early
Clear
Water
Surface
Late
aA sample taken 30 cm from the edge of the pond and the corresponding one taken 1 m from the edge, contained
11
and
6, respectively. The
to
2.
DECEMBER, 1993
pond
very
163
regarded as conservative.
None of these
W.
Ludwig.
1991.
and
95
percent
for 2nd
and
m2
was a
and
invertebrates.
J. Invert.
control
in West
Germany.
mosquito
DECEMBER, 1993
for the same ecological niche. From each water sample, cyanobacteria were characterized on the basis of their
morphological properties using strains previously purified from the same location as references( Thiery et al. 1991).
The highest densities of unicellular cyanobacteria coincided with the peak season of mosquito larvae which usually
extends from April to October. Each water sample was analyzed under laboratory conditions to determine the degree
of ingestion and digestion of unicellular cyanobacteria by larvae of Culex pipiens and Anopheles gambiae
mosquitoes. Although efficiently ingested and digested by mosquito larvae, cyanobacteria did not appear to be
preferentially selected as a food item compared to other microorganisms present in the natural water samples. This
study showed that unicellular cyanobacteria belonging to the genera Synechococcus and Synechocystis are suitable
candidates for mosquito control, in particular those which, according to the criteria used in this work, are identical
to Synechococcus strain PCC 8905 and Synechocystis strain PCC 8906.
characteristics required to colonize certain habitats.
INTRODUCTION
larvae ( Gophen
Khawaled
and
colonize
et al.
larvae
feeding
pipiens(
Bacillus
to control
currently being
but their limited persistence
sphaericus are
used
1983, Davidson
Thiery et al.
et al.
either
B.
were
introduced into
et al.
sphaericus or
B. thuringiensis
var.
israelensis
unicellular cyanobacteria.
recombinant organisms
show
larvicidal
The
activities
laboratory
for
larvae,
the
two
unicellular
strains
more than
in
the physiological
conditions
habitats
However,
have been
mosquito
propagate
1Unit6 des Bacteries Entomopathog8nes, Institut Pasteur, 25 rue du Docteur Roux, 75724 Paris Cedex 15, FRANCE.
2Entente Interdepartementale de DBmoustication, BP 6036, 34030 Montpellier Cedex 1, FRANCE.
3Unit6
rue
DECEMBER, 1993
high larval
season of
constitute a nuisance
the degrees
density during
fed
mosquito species
properties.
digestibility of
algae by two
ingestibility
of
morphological
addition,
cyanobacteria
unicellular
which mosquitoes
165
and
green
and
compared.
Sampling
Methods
From
February
from
immediately
marshland near
withdrawn
of
500
analysed
for
Aliquots
mixed.
ml
were
physico-
and transported
water
and
in
brackish
each of four
were pooled
ml was stored
1991,
February
from
until
and
below).
sterile plastic
samples were
ASN- 10
with
hundred l
medium (
All
Bacto-Agar( Difco).
w/ v)
under
low
photon
flux
density(
plates were
10
Rodier 1966).
vegetation),
as
as
abiotic
parameters ( water
were
percent
of these samples
of each
aliquots
ASN- 10
plated on
3)
serially diluted ( up to 10Thiery et al. 1991). One
incubated
mol/ m2/ s) at
25 C
Cells from
cellular
each
strains)
filamentous
of
or
Feeding Experiment
Twenty-five early 4th- instar larvae of Cx. pipiens
Strains
colony
forming
unit (
filamentous
patches (
CFU)(
or
uni-
according to
according to Rippka
and pigmentation,
et al.
contrast
phase
under
illumination
with
cell
width)
pseudo-
and treated as
described
by Thiery
et al. (
1991).
One
Nikon
Plan 100
properties of the
DL
oil
immersion
cyanobacterial
mosquito
same
Nodar- Lopez
unpub.
strains
logical features
or
were
purified
and
unpub.
that
of
or
from the
by
Thiery
characterized
data),
and
to
the
genus
belonging
Synechocystis
site
Rippka (
Cells
data).
The
previously
breeding
and
Synechococcus
objective.
the
unicellular
strains
determined.
in
CFU
8905"
or
by
counting
all
166
cells
100
for
x(
b-
to a-
1- b/ a). The
The degree
c.
digestion
d) is
equal
was estimated
from
beginning
ingested
the
formula: 100
b-
x( a-
The
cells.
according to
c/ a- b).
DECEMBER, 1993
600
Dynamics
and
in
morphology
identical to those
PCC 8905
or
1991,
and
colonies of
pigmentation
Synechococcus
1990( 800 200 CFU/ ml), had the same size as the latter
of either
Figure 1
Synechococcus"
but was
peak
shows photomicrographs of
type
February to
8905"
density
from 850
varied
season
150
larval
strain
each sample.
CFU/ ml in
of
cyanobacteria whose
were
Abundance
and
Green Algae
was
also
Fig.
recovered
not shown).
Although the
tit
Ail
Figure 1.
'
fin
Yrr15WY
Photomicrographs of the axenic strains Synechococcus PCC 8905( 1) and Synechocystis PCC 8906( 3)
and of the respective homologous strains( 2) and( 4) present in the water samples collected from the
brackish
marshland
located
near
pm).
DECEMBER, 1993
167
1000
Synechococcus" type 8905"( 1 m)
CI
800-
4111k A
600-
400
200
2/ 90
4/ 90
6/ 90
8/ 90
10/ 90
12/ 90
2/ 91
Months
1500 -
Synechococcus"
type
8975"( 13 m)
Synechocystis( 2 m)
Q Synechocystis( 3- 4 m)
a+
gin
1000
41
Synechocystis PE+( 3- 4 m)
t
3
i
500
I.
2/ 90
4/ 90
6/ 90
8/ 90
11166.
10/ 90
12/ 90
2/ 91
Months
Figure 2.
Population dynamics of unicellular cyanobacteria in the brackish marshland located near Fos/mer
France) for the year 1990 with emphasis during the mosquito larva season.( A) Synechococcus strain
type 8905" and Synechocystis strain" type 8906." ( B) Synechococcus strain" type 8975" and newly
found Synechocystis strains. CFU: colony forming units. When calculable, the density variation of
Synechococcus and Synechocystis
was
included between 9- 17
percent and
13- 53
percent,
respectively.
168
DECEMBER, 1993
The
six
different filamentous
strains
were recovered, on at
In
in cell
isolated in
one
addition,
belonging
cyanobacteria
Oscillatoria,( 4- 5
least
latter.
they
new
genus
one
to the
in every
sampling
during
cyanobacteria
the
year and
in
particular
during
For
comparison,
in the brackish
recovered
evaluated(
different
unicellular
TABLE 1).
marshland
At least
green
algae
were
also
digested
cells of
seven
morphologically
distinguished but none of them
were
density
their cell
CFU/ ml
throughout the
they decreased
to only
40 CFU/ ml.
Characteristics
The
brackish
the
Breeding
8905"
and
from 9 C in
These
values were
Precipitation
month, except
in October
139
to
similar
very
meteorological station.
increased
for
winter
October 1990).
logical
Site
seasonal
February
and
of
mm
and
and
nearby
at the meteoro-
to be below 60 mm per
December 1990
112
mm
per
when
it
month,
October 1990
in December 1990.
in Montpellier
Water
and
80 to 100
components
were
cm
analyzed
following
values
indicate the
range
present
in the
water samples(
by Mosquito Larvae.
ingestibility of the different filamentous
strains was recorded after the 60 minutes
feeding of An.
The
gambiae and
Cx.
Most
the strains
except the
PCC 8971 producing phycowhich was never found in the guts of either one
Pseudanabaena
of
larvae.
of
were recovered
erythrin,
pipiens
strain
169
DECEMBER, 1993
Ii.0
3
g."...
e
t
k
m
k $
t
e
Q U
St
U U
c:,
CI)
U
1
t
t
0 \
k /
II
10
CI
2/
k
m
P.
41
7\
E
G.)
A
o
6..
es ]
t $
8
u
2 2
77aa ]
0
2faa /
2 -a
00
t 8/ ]
0
k
0K
c) /
@u
cu
k /
ka
2-
o _%
7..
170
DECEMBER, 1993
water samples
different
be
could
in the
correlated
the abundance
with
because it
allows comparison of
density
of the various
cited organisms.
This study
the
strain
PCC 8905
previously
selected on
Synechocystis
and
larvae,
Thiery
et al.
1991).
be
for
used
Indeed,
strains
light microscopy
200 CFU/ ml
over
be ingested
during
further is
diameter). This
As
interesting
o a
o
M
to study
in
and
in
density between
g Tr CD o
coo co g g
y`
8 8 8
M 8 8
o ao 0
S a
of
Q`
a previous report(
axenic cultures of
Synechococcus
Nov
Qa
c
larvae.
shown
c-
Z1
cell
oo
28
be
large Synechocystis ( 3 to 4
M oR rM N
6 2
and
the larval
digested by mosquito
larvae. Another cyanobacterium found in the course of
season and to
found to be
density
present at a
8 C
with
VD 0000 0000
tin
r"
digestibility by
mosquito control(
a a
a
PCC 8906,
strain
ingestibility
a'
Synechococcus
and
&
a 2)
A a'
tti
3
meaning
o
o
y$
these
of
la
71.
Synechocystis
strain
Thiery et al.
PCC 8905
were
less
likely
and
1991),
PCC 8906
strain
.
o
to be
gNS
v
r
UU
gambiae or
both
Cx.
pipiens
Synechocystis
Synechococcus strain"
show
ingested
in fact
no
strain
distinct
preference.
from 11 to 1
The
size of the
0
cd
m, which corresponds
by
1a
0 0 0
0000
co
co
of
gg&g
8
in the brackish
c,
and
of either An.
8906"
type
"
type 8905,"
larvae
similar quantities of
the
single
dose
of 2.4
'
recombinants
within
24 hours,
whereas,
2.4
killed
larvae
ca.
of
95
Z.,
of Aedes aegypti
the same
Nicolas,
bioassay
are
digested
by
conditions (
in
larvae
release
B.
sphaericus
c", -
`''
personal
mosquito
killed
i
00
percent
Cx. pipiens
g.
a
Ua
a^
E.,
aa
DECEMBER, 1993
Cell
density
in
water sample
Anopheles
Month/ Year
4/90
Culex pipiens
gambiae
CFU/ ml
500
171
CFU/larva
CFU/ larva
800
200
200 _
x259
r 122
1 ^
21540
1540)
6/ 90
300 -
L'
120 ^
1300
7915
120
8/ 90
400
r
I
81';')
1120
4313
2013)
725)
13%
17%
800
160 -
v 4142
763)
, ; :!.
x( 18%
157
1320
1234
i
15100
108
IL
f
1 :"
6%
7%
1120
7137
417)
555)
5717
4*
4/j
2100)
829)
5551
715)
14%
14%
13%
a'
10/90
200
52
150
60
120
6315
2015)
2205
2160
778)
698)
35%
33%
14
r.
32%
12 green- algae
type 8905"
O other cyanobacteria
Figure 3.
Pie diagrams showing the relative cell density of the unicellular cyanobacteria and green algae in four
water samples collected at two-month intervals during the year 1990 compared with the number of cells
ingested per larva after the 60 min. feeding. aTotal number of cyanobacteria and green algae; bnumber
of unicellular cyanobacteria; cthe percentages represent 100 x( b/ a). The numbers of CFU/ml or CFU/
larva of either Synechococcus" type 8905" or Synechocystis" type 8906" are indicated above the arrows
pointing
out the
corresponding
sectors.
172
Culex, but
DECEMBER, 1993
less toxic
are much
Ae.
efficient against
Thiery
and
present
study, An.
REFERENCES CITED
Synechocystis
type 8906"
strain"
during
8905"
and of
the season of
Thus, if
as
Angsuthanasombat, C.
S.
and
Panyim.
1989.
type
8906"
could
brackish
8905,"
be
An.
for mosquito
In contrast,
in the
Chungjatupornchai, W.
1990.
mosquitocidal- protein
Expression of the
genes
of
Bacillus
studied.
since
unpub.
An.
control
B.
marshland
Synechocystis
and
used
likely
be insufficient for
gambiae.
In
conclusion,
Synechocystis
and
strains
belonging
Synechococcus,
to the genera
and
in
particular
type
8905,"
cyanobacterial strains
appear
to
for cloning
be
and
appropriate
expressing the
Abundance of
by
larvae among
Indeed, in
selected
mosquito
microorganisms present
in
water samples.
the
other
Since these
step is the
reintroduction of
cyanobacteria
determine if
maintain
evaluate
in the
and
mosquito
for how
itself in the
breeding
long
mosquito
site.
can
site and to
a given species
breeding
of cells
necessary
to
be
168.
density of the
and
A.
Zaritsky. 1989.
80.
in
collecting
Entente Interdepartementale de Ddmoustication du
samples
and
G. Vigo for
assistance
water
analysis
in the
and to P.
of the cyanobacteria
in
February
1990
field
conditions.
Appl. Environ.
DECEMBER, 1993
M. Herdman,
Generic
assignments,
R. Y. Stanier.
strain
histories
cyanobacteria.
chimique
de l'
Dunod. Paris,
1979.
L' analyse
1966.
Rodier, J.
Waterbury,
173
eau.
Eaux
chimique
et physico-
naturelles- eaux
usees.
412pp.
1593
comparative
H- 14
K. Charoensiri.
persistence of
and
against
toxicity
Bacillus
mosquito
environments.
1983. The
thuringiensis
serotype
of
DECEMBER, 1993
ABSTRACT: The larval development ofAedes aegypti, Anopheles albimanus, Anopheles gambiae, andAnopheles
stephensi were investigated under two different rearing protocols: increasing water depth( 0.5- 14 cm) or increasing
larval densities( 100-> 1500 L/pan) under otherwise constant conditions( 27 C, 14: 10 L:D). The times of pupation
and eclosion were recorded daily, and the protein and lipid reserves were analyzed after eclosion. Defined by the total
eclosion of males and females, the highest larval survival occurred at an optimal water depth of between 0.5 and 1
cm. At greater water depths eclosion was reduced. Anopheles usually did not survive in water deeper than 2 cm,
although dissolved food could extend the maximum depth that would allow survival up to 5 cm for An. albimanus
and An. gambiae. Eclosion of Ae. aegypti was affected less by increasing water levels, in that 50 percent of the
population still eclosed in 14 cm. Larval density strongly affected development, as evidenced by longer duration and
reduced body size. To allow intra- and interspecific comparisons, all caloric data of teneral imagoes were converted
to " size- specific caloric contents."
These values exhibited the synthetic potential for total protein and lipid,
irrespective of actual body size. Aedes aegypti differed from Anopheles. Protein synthesis ofAe. aegypti reached a
fairly constant level at the end of the larval period, regardless of the rearing conditions, whereas the lipid synthesis
increased exponentially when regressed against adult body size. In Anopheles, the corresponding relationships were
consistently on a lower level than in Ae. aegypti despite their overlap in body size. When combining the size- specific
caloric contents of protein and lipid for all larval treatments of Anopheles, their regressions against body size had
identical slopes.
INTRODUCTION
influence
on
primarily vertebrates
Schmidt- Nielsn 1984). The extent of variation within
endocrine
or
developmental
body
size
Insects
for
abnormalities.
several reasons.
First,
their
factors to be
increases
Second,
manipulated.
occur
after
adult
no
eclosion.
by
further
Third,
size
as
In
mosquitoes, the
providing
during
larval
sufficient reserves
is
for
period
responsible
of
Zoology, University
of
function
parameters as a
175
DECEMBER, 1993
of
Briegel 1990a)
aegypti (
and
for four
conditions
extreme
and starvation to
crowding
between
female
body
was
the total
in Ae.
protein of
linearly
were
body
with
in
size
aegypti.
Under
size
body
several culicines,
but in the
body
size did influence the size of the blood meal and therefore
fecundity
the
ingested
ones
of
female Ae.
the
blood
volume of small
eggs
of mature
number
Large females
aegypti.
and
albimanus
showed an exponential
most appropriate (
Anopheles ( Nyssorhynchus)
all
eclosed males
newly
correlated with
increase
size to the
The relationship
close
particularly
females
and
would allow.
and
size
circumstances,
body
allowed us to push
the genome
extremes that
was
almost
et al.
and
quadrupled(
Colless
and
a reason
For the
laboratory
colonization
in
rearing
order to obtain
mosquitoes,
of
homogenous
size(
Lea 1963;
Gerberg 1970;
usually
populations
conditions are
Briegel 1990a,
1990b). This is
physiological
of
1993).
imaginal
a prerequisite
for obtaining
density
The
types in the
identical
under otherwise
efficiency in food
different
acquisition,
imaginal
revealed
such
populations,
mortality in
as
feeding
ultimately
responses
in
already had
of mosquito
1984),
Moore
fecundity (
reflected
of
and
of
development
in terms
assessed
experiments
duration
all stages (
survivorship,
1972; Reisen
and
Previous
body
density- dependent
size.
larval
sense of
meaningful
Barbosaet
oviposition, adult
and
al.
longevity,
and
Hawley 1985;
1991; Kitthawee
performed
to
et al.
1992). Our
measure
et al.
experiments were
individual compensatory
analytical procedures
for caloric
contents were
176
TABLE 1.
DECEMBER, 1993
Feeding regime under standardized conditions for four mosquito species. Spoons
with an average content of 35. 6 0.8 mg and multiples thereof were given in daily
succession. The total amount of the food mixture( mg), added until completion of
development, is converted into caloric input, and expressed per larva.
Aedes
Anopheles
Anopheles
Anopheles
aegypti
albimanus
gambiae
stephensi
Spoon/ day
10
4
4
Total
weights(
Total input(
948
689
1286
544
2938
2137
3987
1687
mg)
cal)
Total caloric
input
per
larva
14.7
7. 1
for individual
caloric values
based
on
5. 6
data
8. 0
to
cal per
body
per
mg
mg lipid. These
size, estimated
by
the
This factor
application
represents an uniform
led to the"
body
size and
its
mosquito."
Two
four
mosquito
water
depth
was
density
In the
kept
identical
for the
series,
until eclosion.
was
an
In the first
hatching
in
species except
RESULTS
manner with
the
cm
to 14
constant
from
larval
per pan to
reaching
DECEMBER, 1993
TABLE 2.
177
Effect of water depth( cm) and two modes of feeding on the percentage of eclosion. Total
females
day
of maximal eclosion(%)
is given.
Modes of feeding
Food
Water
Total
Female
Total
Female
depth
eclosion
eclosion
eclosion
eclosion
day
cm)
Aedes
aegypti
200 larvae)
Anopheles
albimanus
300 larvae)
Anopheles
gambiae
500 larvae)
Anopheles
Food dissolved
sprinkled
stephensi
300 larvae)
day
0.5
49
10
58
89
85
88
75
88
78
82
10
53
93
54
40
11
42
84
11
48
10
59
12
42
76
11
64
14
54
12
48
74
12
41
0. 5
95
10
87
80
11
83
70
11
66
10
17
36
19
16
43
18
42
0. 5
72
10
37
10
38
36
11
57
34
65
13
41
59
14
45
22
15
41
0. 5
89
10
47
92
81
70
11
84
larval
larval
and pupal
development. At
depths in
developmental
females
1 to
a different
Following conventional
all
by only
3 days in Ae.
aegypti.
1 to 2 days in Anopheles
and
added to the
body
was obtained
with
water
water
DECEMBER, 1993
178
depths
of
2 cm,
or
whereas
More informative
lipids,
and carbohydrates.
0. 5
results
were
gained
In newly
cm.
from the
protein,
females the
eclosed
therefore we focused
our
interests
calories and
depths
water level the larval density was varied from 100 larvae
aegypti, grown at
feeding
increased
form
increasing
water
modes.
values
Obviously,
both
success.
was superior.
TABLE 3.
Effect of water depth( cm) with standard densities( larvae/pan) on the caloric content of total protein
P) and lipids( L) of eclosing females of four mosquito species. Means S. E., N= number of females,
at
least
for each
reserve component,
Caloric content
Water Depth
cm)
Ae.
aegypti
200 larvae)
An.
albimanus
300 larvae)
An.
gambiae
500 larvae)
An.
stephensi
300 larvae)
Same
amounts of
P*
L*
E*
0. 5
43
1. 59 0. 15
1. 61 0. 15
0.75 0.27
1. 50 0.29
2. 3
3. 1
93
1. 83 0.20
1. 700. 15
1. 31 0.21
1. 46 0. 32
3. 1
3. 2
93
1. 69 0. 17
1. 71 0. 15
1. 49 0.27
1. 84 0.35
3. 2
3. 5
69
1. 52 0. 15
1. 61 0. 19
1. 05 0.25
1. 85 0.27
2. 6
3. 5
10
56
1. 65 0.23
1. 660. 16
1. 03 0. 22
1. 66 0. 33
2. 7
3. 3
14
52
1. 84 0. 23
1. 56 0.30
1. 23 0.26
1. 75 0. 30
3. 1
3. 3
0. 5
98
1. 12 0. 16
0. 34 0. 10
1. 5
83
1. 14 0. 14
0.45 0. 11
1. 6
70
0.960. 10
0.52 0. 12
1. 5
2*
14
0. 63 0.07
0.29 0.08
3*
23
0.84 0. 10
0. 34 0.09
1. 2
5*
12
0.70 0.28
0. 240. 03
0.9
0. 5
94
0. 63 0. 14
0. 18 0.04
0.8
64
0. 71 0. 10
0. 16 0. 04
0.9
76
0.81 0. 13
0. 16 0. 04
1. 0
2*
170
0.83 0. 13
0.38 0. 19
1. 2
3*
128
0.88 0. 11
0.29 0.07
1. 2
5*
49
0.87 0.07
0.260.05
1. 2
0. 5
125
0.76 0. 12
0. 17 0. 04
0. 9
137
0.67 0. 14
0. 14 0.02
0.8
97
0.75 0. 12
0. 17 0. 04
0.9
ml
distilled
water.
0.9
At
of the
of Ae. aegypti,
densities
all
larval
indicative
Anopheles
density- tolerant
severe reduction
metamorphosis,
TABLE 4). In
percent or more
species(
increasing
as a consequence of
there was a
79
completed
population
of a
179
DECEMBER, 1993
in the
densities,
total number of
larvae
developmental
69
were
11
percent
in An.
from 91
doubled
from 81
percent, and
a reduction
was
these densities
stephensi
percent
once more,
to 59
and to zero
to
caused
If densities
percent.
developmental
in An. stephensi
percent
success
fell to
in An. albimanus.
of
developmental
larval
success
in Ae.
for
the larval
aegypti,
density
An.
was
too
low,
albimanus, or
percent ( TABLE
stephensi.
reduced survival
100 L/ pan
such as
An.
47
density- tolerant
An.
or
reduced
percent.
considered amore
albimanus
than
Alternatively,
when
An.
only
44
to
Therefore,
100 to 1500)
density (
Content
gambiae.
TABLE 4.
of males and
females expressed as
day
percentage(%)
of maximal eclosion(%)
is given.
Eclosion
Larval
density
day
larvae/ pan)
Aedes
1
aegypti
cm)
Anopheles
0. 5
albimanus
cm)
Females
Total
100
79
67
200
93
61
400
82
74
800
81
40
1000
83
11
31
100
81
11
68
300
87
10
50
600
69
13
33
100
73
10
46
500
78
11
51
1000
71
14
29
1500
44
15
28
100
91
55
300
90
10
33
600
59
15
39
1200
11
18
27
11
18
16
1200
Anopheles
0. 5
cm)
Anopheles
gambiae
stephensi
cm)
extremely
crowded
180
by - 56
contents
percent, and -
83
percent
for lipid. In
percent
protein andpercent
for
percent
for
albimanus,-
82
percent
larval lipid
its
percent
synthesis
protein and -
Obviously,
stephensi.
45
the end of the larval period was the same for all species
more
larval synthesis
Methods)
body
size (
increase with body size( Fig. 2). More lipid was deposited
Fig. 1).
For
all
Anopheles
species
per size unit, as a female larva grew larger; this was also
tested,
TABLE 5.
DECEMBER, 1993
larval feeding.
Effect of increasing larval density( larvae/pan) at optimal water levels( cm) on the body size( WL3) of
eclosing females and on the caloric content of total protein ( P) and lipids ( L) at eclosion of four
mosquito species.
reserves,
Means S. E., N=
density
larvae/ pan)
Aedes
1
aegypti
cm)
Anopheles
0. 5
albimanus
cm)
Body
WL3
size
Caloric Content
N
X
3. 4
100
37. 604.40
42
1. 88 0. 10
1. 49 0. 23
200
35. 05 2. 99
95
1. 84 0. 17
1. 32 0.21
3. 2
400
29. 58 3. 69
160
1. 41 0. 21
0. 87 0. 19
2. 3
800
19. 91 4.03
283
0.82 0. 28
0. 35 0. 14
1. 2
1000
21. 53 3. 70
372
0. 97 0.28
0.260.09
1. 2
2. 4
100
46. 53 3. 51
37
1. 760. 15
0.65 0.09
300
34.52 3. 58
130
0.91 0. 10
0. 26 0.05
1. 2
600
24.90 4.75
195
0.49 0. 11
0. 15 0. 02
0.6
1200
0
0
Anopheles
0. 5
Anopheles
1
gambiae
cm)
stephensi
cm)
extremely
crowded
100
34.43 3. 43
41
0.940. 09
0. 48 0. 07
1. 4
500
31. 26 4. 25
138
0.78 0. 14
0.21 0. 06
1. 0
1000
23. 993. 68
171
0.62 0. 18
0. 10 0.02
0. 7
1500
22.29 3. 65
192
0. 52 0. 16
0. 10 0.02
0.6
1. 8
100
36. 24 2. 81
42
1. 17 0. 07
0. 61 0.06
300
27. 77 2. 73
123
0.73 0. 05
0.36 0.05
1. 1
600
21. 87 2. 94
183
0.49 0.09
0.09 0. 02
0.6
1200
20.37 2. 33
66
0.49 0. 10
0. 10 0. 01
0.6
19. 84 2. 68
47
0.42 0.06
0. 11 0.03
0. 5
181
DECEMBER, 1993
0.400
0.300 --
0)
I
w
ig.
11
Go
0.200 --
V
U
V,
U
G.
C/ 1
6
cn
0. 100 --
0-
ti
3. 0
2. 5
50
40
30
20
10
3. 5
I
mm
Body Size
Figure 1.
Size- specific caloric protein( P) and lipid( L) contents per teneral female in relation to body size of all
three Anopheles tested( abscissa: cubic value of wing length above, and linear dimensions in mm below).
Each point is the mean from one experimental cohort of females being raised at variable water depths or
larval densities. Data from TABLES 3 and 5 were recalculated to size- specific caloric contents by using
3),
the formula ( cal x 10 x WL-
Y=0.004X+0. 132( N=21, r=0. 824, p< 0.001) forprotein, and Y=0.004X-0.049( N=21, r=0.777, p<0.001)
for lipids. Total carbohydrates showed no significant correlations with size, therefore only their average
is given( 0.035 0.016, N=21, open square). Symbols are: filled triangles, An. albimanus; filled circles,
An.
gambiae;
filled
squares,
An.
stephensi.
182
0. 600
DECEMBER, 1993
E
0
0.400
a
l
0
U
o
i
U
o
0.200-r
20
30
I
2. 5
20
3. 5
50
1
3. 0
40
'
30
1
2. 5
I
3. 0
40
I
50
1
3. 5
If
mm
Body Size
Figure 2. Size- specific caloric protein( P) and lipid( L) contents per teneral female ofAedes aegypti in relation to
body size( abscissa: cubic value of wing length above and linear dimensions in mm below). Each point
is the mean for one experimental cohort of females being raised at variable water depths or larval
densities, or with different modes of feeding. Data from TABLES 3 and 5 were recalculated to sizespecific caloric contents
by using
the formula (
cal
10
WL-3),
exponential) regression for lipids is Y=0.002X1. 498( N=23, r=0.700). The linear regression for protein
is Y=0.417+ 0. 001X ( N=23,
r= 0.01, n. s.).
and carbohydrates 0.0570.021( N=23, open square). Open circles represent lipid values when food was
dissolved.
In
an attempt to
normalized the
further
eclosed
by
generalize our
data,
we
reserve components)
content (
primarily
synthesis, as
females, therefore
was
calculated
Fig. 3).
per
one
This treatment
of
DECEMBER, 1993
183
Aedes aegypti
2. 000
l
0
r$
1. 000
a.
U
o
o.
10
20
30
40
50
U
U
6
r"
Anopheles
0.600
0.400
0. 200
0
20
1
10
30
1
2. 5
40
1
3. 0
50
1
3. 5
mm
Body Size
Figure 3. Demonstration of increasing larval lipogenesis in relation to protein synthesis, and dependence on body
size for both in Aedes aegypti and three species of Anopheles combined. The size- specific lipid contents
showed in Figure 2, were normalized per unit protein, expressing the lipid per calorie of protein present.
When plotted against body size, a positive allometric correlation was found for Aedes aegypti
Y=0,004X1. 476, N=23, r=0.608) and a linear correlation for the Anopheles( Y=0.011X-0.027, N=21,
r=0.669, p<0. 01). Each point represents the mean of an experimental cohort.
DISCUSSION
potential of mosquito
larvae has
and
primarily
of structural
function,
present
in the
the same
showed
and
lipids:
proportions
in the
synthesis of protein
the
total (
lipids, and
protein.
carbohydrates
1).
Fig.
comparison
had
DECEMBER, 1993
184
and
higher lipid
was a
between larval
findings
other
wing length and protein were equally found in
species ( Van Handel and Day 1989; Briegel, unpubl.
for lipids
Aedes
content
of
result
an
exponential
correlation
our earlier
size,
supporting
Strictly linear correlations between
Briegel 1990a).
in Aedes
also revealed
were
triseriatus(
Briegel,
unpubl. research).
believed to be
ships are
mosquitoes.
of general
field- collected
and
culicine
maintained
validity in
best be
by Day
described
mosquitoes, as
can
relation-
explained
as a
of size,
supply.
In this
These
Differencesbetweenlaboratory-
and
matter
and
albopictus
imagoes. This
proved
allowing the
pression,
results.
have defined
report we
Having
i. e.,
conditions,
findings. Since in
with
represent a
the
larval
In Ae.
content was
level
of
affected
protein
nearly
by
however, the
cal (
to
appear
adult
during
efficiencies.
protein
biomass
identical
size- specific
fairly
and remained at a
Fig.
2).
it
Obviously,
in
was
barely
conditions proceeded
content of
of the
The
larval
size- specific
Anopheles
was all
our
different
and
the
body
lipid
and
1965).
all
Briegel
under
maximal rates.
hypothesis
constant
at
allAe. aegypti
similarity in
the
synthesis
irrespective
to their
the anophelines
size,
acquiring their
higher
0. 42
density
aegypti
environmental
followed the
body
group
metabolic sense,
larval
various
all
content of protein
the
of
relation
comparison
manipulated
water
and
Briegel 1994).
with their
unpubl. research).
were
incapable
of
feeding exclusively
from the
surface
and
ingesting food
consequently depended on
that
explained
few
larvae are
their
by
confined
to shoal biotopes
ability to dive
centimeters,
and
within
can
be
only
dilemma.
energetic
an
suggesting
feed
An.
survival of
An.
albimanus and
gambiae
to
REFERENCES CITED
had
185
DECEMBER, 1993
by
cm
of the teneral
females
Presumably,
the
of
amount
diluted because it
ignoring
our
the increased
volume of water.
surface
certain anophelines.
In Ae.
aegypti
larger
with much
there
They
margins.
could
dive, feed,
and
at
advantage
to the
feeding
to the
domain.
develop
due to
reduced
acquisition might
Anopheles
pools where"
when
the
secondary
level is dras-
water
evaporation.
be crucial,
Dahl
et al. (
1988)
1988)
were unaffected
Anopheles
species appear
by
much
biosynthetic
anophelini,
lower filtration
Culicidae).
in Ae.
482.
ultimately the
the larvae
Acknowledgments
mosquitoes (
appreciate
the reliable
assistance given
generous
support of
by
the
acknowledged.
Linnaeus) ( Diptera:
of culicini and
respectively.
We greatly
aegypti (
and
acquisition
capabilities of
Culicidae).
and yet
larval densities,
to have
and
Rashed
rates then
than Aedes or Culex. These low filtration
efficiencies
filtration rates.
and
species- specific
The latter
rates
every
can
1990a.
resemblance
tically
Briegel, H.
water
crowding"
were
all bionomical parameters tested
notably in
1992.
Nevertheless,
C. M. Holzapfel.
was
results
and
too
per
constant
W. E.
Bradshaw,
container,
food dissolved
held
was
reviewers
critical and
thus helpful
kindly
Diptera: Culicidae).
Mosq.
186
mosquito populations.
Pp. 419-429 in
Ecology of Mosquitoes( Lounibos, L. P., J. R. Rey,
and J. H. Frank, eds.).
Florida Medical Ent. Lab.,
Vero Beach, FL, USA, 579
pp.
mosquito
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ecosystems.
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experimental
rearing and
techniques. Am. Mosq. Contr. Assoc.
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Gillies, G. T. and B. DeMeillon. 1968. The
Anophelinae
of Africa south of the Sahara.
Publ. S. Afr. Inst.
Grimstad, P. R.
and
E. D. Walker.
1991.
and
of
Aedes
La Crosse virus.
larvae
affects the
adult
body size,
R.
Rey,
Florida Medical Ent. Lab.,
J. H. Frank, eds.).
Vero Beach, Fla. USA, 579
pp.
and
Hawley,
W. A. 1985.
The
effect of
and
Aedes
sollicitans.
J.
density
on
955- 964.
Hien, D. S. 1975.
1762)
395.
the
fecundity of Aedes
Culicidae). Ent. Exp. Appl.
on
population.
Klowden, M. J.
parity
J. Med. Ent. 29: 921- 926.
and
Cambridge, 241 p
pp.
biting
Truman, J. W.
and
L. M. Riddiford. 1974.
Physiology
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of insect rhythms.
organization of
the
of
187
young
374- 376.
accumulation
in three
adults.
and J. F. Day.
1989. Correlation
between wing length and protein content of mosquitoes. J. Am. Mosq. Contr. Assoc. 5: 180- 182.
1965.
Van Handel, E.
Wada, Y.
Effect
of
larval
density
on
the
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