Rearing of Texas Tabanidae (Diptera). II. Mass production of Tabanus nigrovittatus Macquart eggs and larvae


Thompson, P.H.; Holmes, J.W.J.; Krauter, P.C.; Raney, C.M.; Clay, M.J.

Southwestern Entomologist 4(3): 224-230

1979


From 60 to 80% of 3000-5000 females of Tabanus nigrovittatus Macq. taken in 5 weekly collections at West Galveston Bay, Texas, in June-July 1978 fed through a lamb-intestine membrane on bovine blood kept at about 100 deg F and ATP (adenosine 5'-triphosphate) was added at concentrations of 10-3 to 10-4 M. Survival, fecundity and fertility were very high in gonotrophically young females taken from this population in late June and early July. About 30 000 newly hatched larvae were obtained from eggs kept in the laboratory under ambient conditions (75 plus or minus 5 deg F and 65 plus or minus 5% RH). Larvae reared on larvae of the stable fly [Stomoxys calcitrans (L.)] in a 2:1 mixture of peat and sand became fully developed in 6-8 weeks but then entered diapause and emerged in April of the following year. Recovery of up to 8 adults per 800 cm3 of substrate suggested that cannibalism was not sufficient to inhibit the rearing of the tabanid en masse. No evidence of mating was observed.

VOL.
4
NO.
3
THE
SOUTHWESTERN
ENTOMOLOGIST
SEPT.
1979
REARING
OF
TEXAS
TABANIDAE
(DIPTERA).
II.
MASS
PRODUCTION
OF
TABANUS
NIGROVITTATUS
MACQUART
EGGS
AND
LARVAE
P.
H.
Thompson,
J.
W.
Holmes,
Jr.,
P.
C.
Krauter,
C.
M.
Raney
and
M.
J.
Clay
Veterinary
Toxicology
and
Entomology
Research
Laboratory
Agr.
Res.
-SEA,
USDA,
College
Station,
TX
77840
ABSTRACT
From
60-80%
of
3,000..5,000
females
of
Tabanus
nigrovittatus
Macquart
taken
on
5
occasions
at
West
Galveston
Bay,
Texas,
fed
on
beef
blood
through
prophylactic,
when
the
blood
was
held
at
ca.
100°
F
and
ATP
was
added
at
10'
to
10
-
4
M.
Survivorship,
fecundity
and
fertility
were
very
high
in
gonotrophically
young
females
taken
from
this
population
in
late
June
and
early
July.
Approximately
30,000
emergent
T.
nigrovittatus
larvae
were
obtained
from
eggs
held
in
the
laboratory
under
ambient
conditions
(75
°
F±5
and
65%.±5
R.H.).
Larvae
reared
on
stable
fly
maggots
in
a
2:1
peat
-sand
mixture
matured
in
6-8
weeks
but
then
entered
diapause
and
emerged
in
April
of
the
following
year.
Recovery
of
up
to
8
adults
per
800
cc
of
substratum
suggested
that
cannibalism
did
not
deter
rearing
specimens
en
masse.
No
evidence
of
mating
was
observed.
RES1JMEN
Entre
60-80%
de
3,000-5,000
hembras
de
Tabanus
nigrovittatus
Macquart
fueron
coleccionadas
5
veces
en
la
Itahia
de
Galveston,
Tejas
y
fueron
alimentadas
sangre
de
res
por
membrana
intestinal
de
oveja
cuando
la
sangre
fue
wanteniaa
aproximadamente
a
100°F
y
ATP
fue
agregado
a
razdn
de
10
-
-'
a
10
-4-
M.
Supervivencia,
fecundidad,,
y
fertilidad
fueron
muy
altar
en
hembras
jovenes
de
esta
populacion.
A
proximadamente
30,000
larvas
emergentes
de
T.
nigrovittatus
fueron
obtenidas
de
huevecillos
mantenidos
en
el
laboraforIo
a
condiciones
ambientes
(75
°
F!5
e
65%1:5
R.H.).
Larvas
de
esta
especies
criadas
al
alimentarlas
con
larvas
de
la
mosca
del
establo
en
un
substrato
de
2
partes
dl
turba
por
una
de
arena,
se
desarrollaron
en
6-8
semanas
pero
despues
entraron
diapausa
e
emergieron
en
abril
del
ano
siguiente.
No
se
observo
s
evidencia
de
canibalismo.
No
se
observo'
evidencia
de
conducta
reproductivo.
INTRODUCTION
Thompson
and
Krauter
(1978)
fed
large
quantities
of
Tabanus
nigrovittatus
Macquart
females,
and
smaller
numbers
of
T.
lineola
var.
hinellus
Philip
and
T.
acutus
(Bigot),
on
warmed,
citrated
beef
blood
held
in
prophylactics
derived
from
lamb
intestine
and
placed
on
top
of
trap
collection
cages.
These
females
were
subsequently
held
at
ambient
temperatures
and
humidities
in
the
laboratory
to
obtain
eggs
for
rearing
and
colonization
attempts.
The
following
report
describes
the
results
of
these
attempts
made
during
the
summers
of
1977
and
1978.
224
=HODS
AND
RESULTS
Collection
of
Flies.-
Five
weekly
collections
of'Tabanus
nigrovittatus
were
made
from
June
20
through
July
25,
1978,
using
Animal
Traps
(ATs)
baited
with
dry
ice.
Trapping
was
begun
in
midafternoon
and
continued
through
dusk
at
which
time
catches
were
transported
to
the
laboratory
in
insulated
picnic
chests
chilled
with
liquid
refrigerant.
Retention
in
the
Laboratory.-
Female
flies
were
initially
held
in
half-pint
ice
cream
cartons
in
the
laboratory
but
no
eggs
were
deposited
by
2-200
engorged
specimens
representing
the
3
salt
-marsh
species
collected
in
this
study
and
T.
abactor
Philip
and
T.
mularis
Stone.
Subsequent
examination
of
thg
oWalgETollicles
of
-
thg
-5
----
species
showed
that
the
ovaries
had
remained
in
a
state
of
ovarian
arrest
although
no
less
than
7
days
had
expired
since
the
insects
engorged.
Subsequent
attempts
to
obtain
eggs,
therefore,
were
conducted
while
holding
females
in
AT
cages
(1(a.
in.
X
9.5
in.
X
9.5
in.).
The
design
of
the
collection
container
was
converted
as
a
laboratory
cage
in
the
following
manner.
The
cellulose
acetate
funnel
serving
as
a
baffle,
which
was
initially
stapled
to
the
bottom
of
the
collection
container
(Thompson
1969),
was
replaced
with
one
of
plastic
window
screen
now
stapled
to
the
trap.
As
a
result
of
this
change,
the
floor
of
the
cage
was
formed
not
by
this
baffle
but
by
a
movable
sheet
of
galvanized
metal
flashing,
held
beneath
flanges
of
the
same
material
nailed
to
the
bottom
of
the
wood
cage
frame.
After
the
container
accumulated
250-300
females,
these
2
modifications
enabled
the
collector
to
enter
the
cage
through
the
stockinette
sleeve
in
one
end,
depress
the
plastic
screen
funnel
down
within
the
trap
body,
and
close
the
metal
floor
of
the
cage.
Approximately
3,000-5,000
females
were
taken
in
5-6
hrs
of
collecting
during
the
first
4
weeks
of
trapping.
Catches
of
the
last
week
(July
25)
suggested
that
this
T.
nigrovittatus
population
was
beginning
to
decline
at
that
time.
This
decline
was
associated
with
an
increase
in
parity
and
with
a
striking
decrease
in
fecundity,
fertility,
and
larval
vigor.
Anticoagulation
vs.
Defibrination.-
High
mortalities
of
fed
flies
during
the
first,
and
preceding,
summer
of
study
(1977)
indicated
that
sodium
citrate
was
toxic.
To
resolve
this
problem,
flies
were
fed
on
defibrinated
beef
blood
during
1978
and
no
additives
of
any
kind
were
introduced
thereafter
(with
feeding
stimulant
accepted).
Survivorship
of
fed
flies
markedly
increased;
i.e.,
the
five-day
survivorship
of
the
1977
sample
of
410
flies
fed
on
citrated
blood
was
48.5%
(Thompson
and
Krauter
1978)
whilethat
of
the
1978
sample
of
600
flies
fed
on
defibrinated
blood
was
76.7%.
1/
Feeding
Stimulants.-
After
Fourex
Natural
Skins
(Schmid
Laboratories,
Inc.,
Little
Falls,
NY
07424)
were
rinsed
with
hot
water
to
remove
additives,
they
were
filled
with
20
ml
of
blood
-
rather
than
the
30
ml
used
previously
-
so
that
the
condoms
would
flatten
over
a
larger
area
of
the
screened
cage
ceiling
during
feeding.
The
blood
was
introduced
and
then
adenosine
s
5I-triphosphate
or
ATP
(Sigma
Chem.
Co.,
St.
Louis,
MO
63178)
-'was
1/
This
paper
reports
the
results
of
research
only.
Mention
of
a
commercial
product
here,
and
elsewhere
in
this
publication,
does
not
constitute
a
recommendation
for
use
by
the
United
States
Department
of
Agriculture.
225
added
at
10
-
3-10
-
4
M.
Initial
results
with
these
ATP
concentrations
were
so
successful
that
individual
portions
were
only
measured
with
a
spatula
thereafter.
Temperature.-
Wide
variation
in
early
feeding
results
with
ATP
suggested
that
temperature
was
a
critical
stimulus
in
feeding
this
horse
fly
species.
Therefore,
because
of
the
avidity
of
T.
nigrovittatus
for
cattle,
blood-filled
prophylactics
were
heated
To
approximate
the
normal
body
temperature
of
a
cow.
After
the
condoms
were
initially
immersed
in
warm
water,
they
were
maintained
at
100
0
F
using
a
microscopic
slide
warmer.
Sufficient
prophylactics
were
then
prepared
to
cover
most
of
the
ceiling
surface
of
the
cage
(usually
12)
and
then
the
inverted
slide
warmer
was
placed
over
them,
separated
by
a
polyethylene
bag
to
prevent
the
membranes
from
sticking
to
the
plate
of
the
warmer.
The
weight
of
the
warmer
was
supported
by
wooden
frames,
placed
at
either
side
of
the
fly
cage,
so
that
the
warmer
plate
only
depressed
the
blood-filled
containers
sufficiently
to
warm
their
contents.
Feeding
markedly
increased
from
an
estimated
5-30%
to
an
estimated
60-80%
using
this
procedure.
After
the
prophylactics
were
placed
atop
a
cage
of
unfed
flies,
the
insects
became
voracious
and
either
penetrated
the
membranes
immediately
or
flew
to
them
to
find
a
place
to
feed.
Feeding
was
usually
completed
within
a
minute
and
no
females
remained
feeding
within
5
minutes
after
exposure;
the
probing,
sampling,
and
engorgement
phases
of
feeding,
so
often
apparent
as
separate,
sequential
responses
during
less
successful
exposures
without
ATP
or
constant
temperature
control,
were
inapparent
here.
Reuse
of
Membranes.-
Prophylactics
were
used
3-4
times
without
any
apparent
effect
on
feeding
success
or
fecundity.
They
were
rinsed
after
use
and
then
soaked
in
Septisol
(Vestal
Labs.,
St.
Louis,
MO
63110)
for
an
hr
or
so;
then
thoroughly
rinsed
again
and
frozen
wet'until
use
again.
In
addition,
these
membranes
were
resilient
enough
to
withstand
many
feeding
punctures
without
leaking
to
the
point
of
impeding
engorgement.
Oviposition.-
Tabanids
laid
eggs
under
laboratory
conditions
of
75°Ft5,
6505
R.H.,
and
outdoor
lighting
from
one
large
wall
window.
Females
of
T.
nigrovittatus
were
seen
to
oviposit
during
midday
on
5
occasions;
This
species
laid
eggs
primarily
on
wood
cage
frames
and
on
the
moist
surface
of
red
clay
flower
pots
holding
grasses
whereas
T.
lineola
var.
hinellus
laid
eggs
on
the
grasses
themeelvgs.
Egg
Characteristics.-
The
eggs
of
T.
nigrovittatus
and
T.
lineola
Ilinellus
were
usually
deposited
in
a
tiers
of
long
columns,
while
those
of
T.
acutus
were
shingled,
rather
than
tiered,
in
a
triangular
mass":
TEZRgs
of
these
3
spp.
were
easily
separated
using
the
color
and
surface
characters
defined
in
Table
1.
Eggs
of
T.
nigrovittatus
were
covered
with
an
opaque,
amorphous
coating
wile
hose
or
the
other
2
spp.
were
not.
In
addition,
those
of
T.
lineola
var.
hinellus
were
uncoated
and
translucent,
so
that
as
tHe
larvae
completed
embryonic
development,
segmentation
and
movement
were
apparent
through
the
chorion.
TABLE
1.-
Size,
Form,
and
Coloration
of
the
Eggs
of
3
Species
of
Coastal
Marsh
Tabanids
(Samples
of
10
Eggs
Each
from
3
Females).
Species
Width
(mmi
Length
(m)
Wit
Ratio
Color
Cnorion
coat
T.
T.
T.
acutus
0.3
0.2
0.29
2.43
1.33
1.67
0.152
white white
0.210
clear
none
0.173
brown
white
lineola
var.hinellus
7117(Wittatus
226
Harvesting
the
Eggs.-
The
maximum
yield
of
T.
nigrovittatus
eggs
was
produced
by
females
taken
during
the
median
week
of
collection
(July
5).
After
females
began
to
oviposit,
the
peak
occurring
at
8-9
days
after
feeding
in
the
laboratory,
eggs
were
harvested
every
3-4
days.
The
maximum
3
-day
yield
of
eggs
laid
by
July
5
females
filled
a
30
-cc
petri
dish.
Early
lots
showed
no
mold
growth
and
only
rarely
did
a
mass
fail
to
hatch.
Care
of
the
Masses.-
The
first
(1977)
attempts
to
increase
atmospheric
moisture,
and
thereby
prevent
dessication
of
eggs,
were
unsatisfactory
because
most
became
infected
with
molds.
However,
experience
during
1978
showed
that
existing
ambient
humidities
in
the
laboratory
(6505
R.H.)
were
ideal
for
maintaining
viable
egg
development.
Eggs
were
recovered
from
cage
and
plant
surfaces
by
sliding
the
upper
rim
of
a
small
cone,
fashioned
from
5.5
cm
filter
paper,
between
the
egg
mass
and
the
substratum.
These
harvested
masses
were
held
in
receptacles
made
from
the
rims
of
Mason
Jars,
the
lids
of
which
were
replaced
with
circular
sections
of
60
-mesh
wire
screen
(Brass
Strainer
Cloth,
C.
O.
Jelliff
Corp.,
Southport,
CT
06490).
The
mesh
of
this
screen
retained
developing
eggs
while
allowing
the
emerging
larvae
to
drop
down
to
the
rearing
medium
beneath.
Egg
receptacles
containing
not
more
than
2
layers
of
eggs
were
held
over
the
moistened
rearing
medium
until
the
time
of
hatching
approached.
Stimulating
Eclosion.-
The
habit
of
tabanid
larvae
remaining
within
the
egg
after
they
have
completed
development
was
utilized
in
controlling
hatch,
and
thus
managing
larval
densities
in
rearing
trays.
(Lecaillon
1905,
after
Marchand
1920,
attributed
the
widely
varing
incubation
periods
of
Tabanus
quatuornotatus
Meigen
to
this
mechanism.)
Eggs
contained
matilF17376nryos
at
6-7
days
after
deposition
in
our
laboratory
at
75°F±5.
To
provoke
the
larvae
to
emerge,
the
screened
Mason
Jar
lid
containing
eggs
was
moved
from
the
surface
of
the
rearing
medium
to
the
surface
of
a
9
-cm
glass
petri
dish
holding
a
sheet
of filter
paper.
The
edge
of
this
lid
was
then
touched
with
a
vibrator
-massager
(Deep
Heat
Massager,
Montgomery
Ward,
St.
Louis,
MO
60607),
causing
the
larvae
which
were
then
ready
to
emerge
to
do
so
en
masse.
The
edge
of
the
filter
paper
was
tapped
over
the
tray
of
rearing
medium
until
ca.
100
larvae
had
dropped
into
it.
However,
the
surface
of
the
filter
paper
was
checked
between
such
2
-hr
intervals
in
order
to
prevent
young
larvae
from
drying
out.
For
this
reason,
too,
the
egg
receptacles
were
transferred
to
the
surface
of
the
rearing
medium
before
we
left
the
laboratory
for
the
evening.
Substrata.-
Throughout
their
development,
larvae
were
maintained
in
a
2:1
mixture
of
sphagnum
peat
and
washed
sand.
To
prevent
these
materials
from
separating,
both
were
sifted
before
admixture,
the
sand
through
42
-mesh
screen
and
the
peat
through
24
-mesh
screen.
This
medium
was
placed
in
1
-gal
plastic
ice
cream
pails
at
ca.
800
cc
per
container
and
saturated
with
distilled
water
and
live
maggots
of
stable
flies
were
provided
as
food
3
days
weekly.
The
mouths
of
the
pails
were
covered
with
a
square
of
12
-mesh
organdy
which
was
held
in
place
with
a
rubber
band.
Late
2nd
and
early
3rd
instar
maggots
were
separated
by
salt
flotation
from
their
rearing
medium,
skimmed
from
the
surface
with
a
strainer,
and
transferred
to
the
organdy
pail
cover
where
they
wiggled
through
the
holes
in
the
material
and
dropped
to
the
substratum
beneath.
Development
and
Emergence.-
Larvae
matured
in
6-8
weeks
but
then
entered
diapause
before
pupation
and
did
not
emerge
as
adults
until
the
following
spring.
Emergence
began
the
last
week
of
April,
peaked
with
32
specimens
in
the
3rd
week
of
June,
and
dwindled
to
227
1-3
specimens
per
week
throughout
September.
Fifty-nine
of
81
pails
maintained
under
the
ambient
laboratory
conditions
previously
described,
yielded
212
adults
(107
females
and
105
males).
Evidence
Against
Cannibalism.-
Recovery
of
multiple
adults
from
most
of
these
59
pails
showed
that
cannibalism
did
not
deter
rearing
en
mass
(Table
2).
In
addition,
the
examination
of
moribund
specimens
showed
no
such
evidence,
i.e.
no
eviscerated
bodies
or
torn
integuments.
TABLE
2.-
Frequency
Distribution
of
T.
nigrovittatus
Adults
Successfully
Reared
from
59
Pails,
Each
Containing
ca.
800
cc
of
Substratum.
Number
of
Larvae
1
2
Number
of
Pails
9
10
6
7
8
0
Reproductive
Behavior.-
Adult
insects
were
freed
into
a
screened
cage
constructed
from
2
in.
X
4
in.
-framing,
the
outside
dimensions
of
which
were
2
ft
X
2
ft
X
4
ft.
This
cage
was
placed
beside
a
window
in
the
laboratory
and
the
insects
were
provided
with
a
10%
aqueous
honey
solution
daily.
Survivorship
was
high
and
30
or
more
adults
were
found
in
the
cage
at
one
time
during
peak
emergence
in
late
June.
There
was
no
evidence
of
swarming
or
sexual
activity
during
the
day
or
at
dusk
or
dawn.
No
eggs
were
found
in
the
cage
and
no
sperm
were
found
in a
sample
from
this
population.
The
ovaries
of
these
25
females
showed
no
ovarian
development
beyond
Nor's
Stage
I
-II
(Mer
1936).
DISCUSSION
AND
CONCLUSIONS
Information
on
the
rearing
and
development
of
Tabanus
nigrovittatus
and
T.
lineola
var.
hinellus
is
virtui717Taexistent.
Small
numbers
of
large
f
rvae
and
pupae,
collected
from
the
field,
have
been
reared
in
conjunction
with
survey
and
taxonomic
study
by
several
workers
but
no
reports
have
been
published
on
the
en
masse
rearing
from
eggs
which
we
have
reported
here.
Also,
no
observations
on
the
reproductive
habits
of
T.
lineola
var.
hinellus
have
been
reported
although
such
observations
have
concerned
similar
salt
-marsh
forms
of
the
northeast.
Teskey
(1969)
examined
material
studied
by
Jamnback
and
Wall
(1959)
and
concluded
that
they
were
dealing
with
what
he
called
the
"northern
salt
marsh
form"
of
T.
lineola
F.
but
the
identity
of
the
material
studied
by
OrmTnati
and
Hansens
(1974)
is
questionable.
Both
Jamnback
and
Wall
and
Orminati
and
Hansens
found
large
numbers
of
egg
masses
on
Spartina
and
Distichlis
-
masses
which
probably
represent
Teskey►s
northern
salt
-marsh
form
of
T.
lineola
in
both
cases.
Our
encouraging
results
with
T.
nigrovittatus
have
depended
upon
the
abundance
of
this
horse
fly
in
-
the
marshes
of
West
Galveston
Bay
where
collection
of
females
by
the
thousands
in
a
few
hours
is
not
a
difficult
task;
and
where
this
species
is
apparently
unaffected
by
the
weather
conditions
decimating
local
populations
of
abundant
inland
species
(Thompson
1977).
Survey
data
showed
that
the
most
efficient
yields
of
physiologically
young
females
from
this
population
will
be
taken
from
late
June
through
early
July.
Although
our
initial
studies
with
T.
lineola
var.
hinellus
and
T.
acutus
indicate
these
forms
have
rearing
potential,
their
smaller
numbers
are
extremely
vulnerable
to
the
selection
pressures
inherent
in
rearing
attempts.
228
Temperature
and
chemical
stimulation,
in
addition
to
host
blood
source
and
membrane
composition,
are
factors
which
greatly
improved
engorgement
and
egg
production.
Also,
the
use
of
sodium
citrate
as
an
anticoagulant,
a
usage
we
suspected
of
causing
mortality
to
horse
flies
in
our
work,
was
reported
to
do
so
in
tsetse
flies
(Laird
1977).
We
separated
the
eggs
of
the
3
spp.
taken
at
West
Bay
using
size,
color,
and
surface
character
and
thereby
obviated
separation
of
females
by
species
after
their
collection.
But
such
separations
should
be
made
with
caution
because
eggs
show
the
magnitude
of
differences
described
for
adults
of
widely
-separated
geographic
populations.
The
T.
nigrovittatus
egg
masses
described
by
Jamnback
and
Wall
(1959)
were
similar
to
those
which
we
observed
on
vegetation
-
elongate,
linear,
narrowed
apically,
and
usually
consisting
of
2
tiers
and
several
rows
but
the
individual
eggs
were
longer,
more
heavily
bodied,
and
apparently
lacked
a
coating
or
dense
pigmentation.
Tabanus
lineola
var.
hinellus
was
described
by
Philip
(1960)
from
material
collected
in
eastern
Louisiana
and
its
range
extends
to
Alabama
(Watson
1968)
but
the
status
of
related
coastal
marsh
forms
from
other
regions
of
the
Atlantic
Coastal
Plain
remains
undefined.
They
do,
however,
also
differ
from
populations
of
the
darker
inland
form,
T.
lineola
F.,
which
predominates
on
river
floodplains
throughout
the
eastern
U.S.
(Fairchild
1942,
Philip
1942,
and
Teskey
1969).
Comparative
differences
concerning
the
eggs
and
development
of
these
species
should
be
considered
with
this
taxonomic
background
in
mind.
Other
techniques
and
materials
were
also
improved
for
handling
and
transporting
catches;
and
for
harvesting
eggs,
stimulating
eclosion,
and
managing
larval
density.
However,
our
recent
experience
with
Tabanus
nigrovittatus
suggests
that
several
obstacles
to
colonization
remain:
to
prevent
diapause
of
larvae
and
to
induce
mating
of
adults.
ACKNOWLEDGMENT
We
gratefully
acknowledge
the
help
of
Carolyn
Love,
Theresa
Holland,
James
Linklater,
and
Tatiana
Araujo
with
rearing;
of
R.
L.
Harris,
Alan
Bridges,
Ellen
Steele,
B.
F.
Hogan,
R. T.
Mayer,
J.
L.
Durrant,
S.
M.
Meola,
J.
M.
Thompson,
Ivan
Berry,
F. E.
Wood,
and
J.
0.
Norman
with
various
procedures
and
materials;
and
of
L.
L.
Pechuman,
H.
J.
Teskey,
S.
M.
Meola,
C.
Sheppard,
and
A.
B.
Bosworth
with
constructive
criticism
of
the
manuscript.
LITERATURE
CITED
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G.
B.
1942.
Notes
on
the
Tabanidae
(Diptera)
from
Panama.
VII.
The
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Amer.
35:
153-182.
Jamnback,
H.
and
W.
Wall.
1959.
The
common
salt
-marsh
Tabanidae
of
Long
Island,
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Y.
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No.
375,
77
pp.
Laird,
M.
1977.
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the
future
of
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v.
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Ctr.,
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220
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Lecaillon,
A.
1905.
Sur
la
ponte
des
oeufs
et
la
vie
larvaire
des
Tabanides,
particulierment
du
Taon
a
quatre
taches
(Tabanus
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74:
20-28.
Marchand,
W.
1920.
The
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No.
13,
Rockefeller
Inst.
Med.
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203
pp,
15
pl.
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G.
G.
1936.
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ovary
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A.
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lineola
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---
Philf57T.
--
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1942.
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Part
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The
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lineola
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49:
25-40.
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1960.
New
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Supplemental
notes
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to
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catalog
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species.
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53:
364-369.
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H.
J.
1969.
Larvae
and
pupae
of
some
eastern
North
American
Tabanidae
(Diptera).
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Entomol.
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63,
147
pp.
Thompson,
P.
H.
1969.
Collecting
methods
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Soc.
Amer.
62:
50-57:
Thompson,
P.
H.
1977.
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(Diptera)
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Texas.
VII.
Comparisons
of
upland
and
lowland
tabanid
populations
in
southeast
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79:
564-574.
Thompson,
P.
H.
and
P.
C.
Krauter.
1978.
Rearing
of
Texas
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(Diptera).
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Collection,
feeding,
and
maintenance
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marsh
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80:
616-625.
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R.
L.
1968.
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