Medical and Veterinary Entomology (2006) 20, 11–26

Orientation of Culex mosquitoes to carbon dioxide-

baited traps: flight manoeuvres and trapping efficiency
M . F . C O O P E R B A N D and R . T . C A R D É
Department of Entomology, University of California, Riverside, U.S.A.

Abstract. Females of Culex quinquefasciatus Say and Culex tarsalis Coquillet

(Diptera: Culicidae) in the host-seeking stage were released and video recorded
in three dimensions in a large field wind tunnel as they flew to four kinds of
CO2-baited mosquito traps. The trapping efficiency (number of mosquitoes
approaching compared to the number caught) was determined for each trap
type. The Encephalitis Virus Surveillance (EVS), Mosquito Magnet Freedom
(MMF) and Mosquito Magnet Liberty (MML) traps captured only 13–16% of
approaching Cx. quinquefasciatus females, whereas the Mosquito Magnet-X
(MMX) trap captured 58%. Similar results were obtained for Cx. tarsalis.
Orientation behaviour and flight parameters of mosquitoes approaching the
four traps were compared. Mosquitoes spent the most time orienting to the
EVS trap. Flight speed decreased as mosquitoes entered the vicinity of each
trap and a large portion of their time was spent within 30 cm downwind of the
traps. Flights became highly tortuous downwind of the poorly performing traps
and just upwind of the MMX trap. Differences between traps and possible
explanations for the superior performance of the MMX trap are considered.
Key words. Culex quinquefasciatus, Culex tarsalis, anemotaxis, carbon dioxide-
baited traps, host odours, trapping efficiency.

Introduction plume (Dekker et al., 2001). Dekker et al. (2005) also

Female mosquitoes detect changes in CO2 concentration as
minute as 50 p.p.m. via sensilla on their maxillary palps
(Grant & O’Connell, 1996). When a host-seeking female
encounters a plume of CO2, she orients upwind using opto-
motor anemotaxis (Kennedy, 1939; Daykin et al., 1965).
The structure of the plumes of CO2 and other host-odour
kairomones plays an important role in the attraction of
Aedes aegypti (L.) mosquitoes. In a Y-tube olfactometer,
orientation behaviour of Ae. aegypti varied with plume
structure and odour, with a filamentous presentation of
CO2 inducing improved upwind movement over a homo-
genous cloud of CO2 (Geier et al., 1999). In a dual choice
flight chamber, fewer Ae. aegypti females entered ports
emitting a homogenous CO2 plume than a turbulent
plume, whereas more females entered ports emitting a
homogenous plume of human skin odour than a turbulent
found that females were induced in a wind-tunnel assay to
surge upwind by entering a concentration of CO2 as little as
0.5% above ambient, and that following a brief encounter
with a single filament of CO2, mosquitoes were sensitized to
fly upwind to lower concentrations of human skin odour
than without such prior CO2 exposure.
Public health and vector control programs designed to
predict and control disease epidemics rely in part upon
early detection of infected mosquitoes. This requires sam-
pling of mosquito populations to detect population surges,
delimit their distribution and establish the presence of the
infected mosquitoes. Important tools in such monitoring
programs are CO2-baited mosquito traps (Kline, 1999;
Rueda et al., 2001; reviewed in Service, 1993). These traps
have also been suggested as possible tools in mosquito
control (Curtis, 1996). The release of CO2 by these traps
attracts female mosquitoes in the host-seeking stage.

Correspondence: Miriam F. Cooperband, Entomology, University of California, Riverside, CA 92521, U.S.A. Tel.: þ 1 951 8274492;
fax: þ 1 951 8273681; e-mail: mcooperband@alumni.ucr.edu

# 2006 The Authors

Journal compilation # 2006 The Royal Entomological Society 11
12 M. F. Cooperband and R. T. Carde´
Adding 1-octen-3-ol (octenol), an odour from ox breath to
which tsetse flies are attracted (Vale & Hall, 1985; Takken &
Kline, 1989; Kline et al., 1990; Kemme et al., 1993; Kline,
1994), or heat (Kline & Lemire, 1995; Makiya & Iwao, 2001),
can improve trap capture for some mosquito species.
CO2-baited traps have been compared under a wide vari-
ety of field conditions to evaluate the roles various configu-
rations and odour release characteristics could play on
trap capture (Hayes et al., 1958; Reisen & Pfuntner, 1987;
Cummings & Meyer, 1999; Kline, 1999, 2002; Mboera et al.,
2000; Reisen et al., 2000; Burkett et al., 2001). These studies
generally have compared the numbers of different species of
mosquitoes captured by different trap types using various
combinations of visual or chemical cues such as light, CO2
and octenol. Because most of these studies were carried out
in the field, there is no information on the trapping effi-
ciency, that is, what proportion of mosquitoes lured to the
vicinity of a given trap is actually captured.
Vale & Hargrove (1979) addressed the question of field
trapping efficiency for tsetse flies using electric nets (pre-
sumed invisible to the flies) strategically placed around a
given trap configuration. This innovative method enabled
comparison of the number of flies arriving in the vicinity of
the trap with the number actually captured. Studies to
determine trap efficiency with mosquitoes have released
set numbers into a large, outdoor cage, in which a trap
was operated from 90 min before dusk to 90 min after
dawn (Kline, 1999, 2002). Because they deal with a constant
number of mosquitoes, they provide useful comparisons of
the relative success of traps. Such studies, however, cannot
estimate trapping efficiency, because a mosquito enclosed
within the cage for approximately 12 h will have multiple
opportunities to approach a trap, whereas in the field a
mosquito that approaches a trap but is not caught may
subsequently leave its vicinity. In such instances, the reason
approaching mosquitoes are not captured would be
unknown. Trapping efficiency can be determined by
examining the proportion of mosquitoes approaching
traps that are actually caught. In our previous study
(Cooperband & Cardé, 2006), we examined the differences
between trap designs, CO2 plumes and other potential cues
produced by the same four traps. In this study,
Cx. quinquefasciatus and Cx. tarsalis are used to compare
trapping efficiency and to examine the flight manoeuvres of
female mosquitoes as they orientate upwind along the CO2
plumes emanating from these traps.
Materials and methods
Wind tunnel
Experiments were conducted in a field wind tunnel
(Cooperband & Cardé, 2006), similar to the ‘pushing’
wind tunnel described by Cardé et al. (1998). The steel
frame of the wind tunnel was 2.5 m wide
1.85 m high

6.2 m long, and was covered with a translucent polyethy-
lene plastic sheet which formed the ceiling and walls and
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
had vertical zipper doorways on the sides near the middle
and the upwind end. Air entering and exiting the wind
tunnel passed through two screens made of HexcelÒ
(aluminium sheet of hexagonal cells 5 cm thick with 1 cm
I.D.) covered with black nylon insect screening to reduce
turbulence. A fan (60 cm diameter blades) on a metal stand
0.8 m above the ground mixed and pushed outside air
through the tunnel via a polyethylene pre-chamber which
expanded over a distance of 2.5 m from the fan to the
upwind laminizer. Except for the frame, which was fixed
in the ground, the wind tunnel was assembled before each
evening of experiments and at the end of experiments was
dismantled and stored. Latex gloves were worn while set-
ting up, running experiments and dismantling, and care was
taken to avoid handling with bare skin anything to which
mosquitoes would be exposed. Prior to set-up, the earth
floor of the wind tunnel was cleared of any plants, twigs,
rocks, leaves or other obvious visual cues to ensure similar
conditions from day to day. Other large visual cues that
would allow optomotor anemotaxis to take place remained,
such as the wind tunnel frame, the video camera and tripod,
the IR lighting, the laminizer screens, large stationary
objects outside of the wind tunnel, and the traps them-
selves. A hot wire anemometer was used to measure the
wind speed, which was adjusted to 50 cm/s.
Traps
Four CO2-baited mosquito traps (see Cooperband &
Cardé, 2006) were compared: encephalitis virus surveillance
(EVS) trap, Mosquito MagnetÒ Freedom (MMF),
Mosquito MagnetÒ Liberty (MML) and the Mosquito
MagnetÒ X (MMX).
The EVS trap consisted of an insulated plastic bucket
(17 cm diameter by 20 cm high) to house dry ice with four
0.5 cm holes about 1 cm from the bottom to release subli-
mated CO2 (Rohe & Fall, 1979). Hanging below the bucket,
three 1.5 V batteries powering the trap fan rested on a
circular disc, attached 16 mm above the opening of a cylin-
der (9.5 cm high
11.5 cm diameter) which housed the fan.
The battery-operated fan pulled mosquitoes through the 16-
mm opening into a net at the bottom of the cylinder. The
CO2 outlet holes on the bucket were 20 cm above the trap
entrance and the bucket contained 1.8 kg of dry ice.
The MMF and MML were designed by American
Biophysics Corp. (ABC) (North Kingstown, RI, U.S.A.).
They utilize two fans, or ‘counter-flow technology’, in
which one fan exhausts odours out the bottom outlet,
while the other fan draws air from the bottom inlet and
forces it out at the top, causing a suction at the bottom inlet
that pulls in mosquitoes. Both traps rest on a stand on the
ground and run on propane, which produces energy for the
fans and releases CO2, heat and humidity as a by-product
of combustion. The release of heat has been suggested as a
way to improve trap efficacy (Kline & Lemire, 1995;
Makiya & Iwao, 2001). The CO2 outlet is 10 cm below
the trap entrance in all the Mosquito MagnetÒ traps. In
# 2006 The Authors

the MMF and MML, heated, humidified CO2 is pushed
out through that outlet. These traps differ slightly in size,
shape, inlet and outlet airflow, CO2 concentration, heat,
humidity and plume structure (see Cooperband &
Cardé, 2006).
The MMX counterflow geometry trap was designed by
ABC for research purposes and is not available commer-
cially. It was first described by Kline (1999) and was found
to capture more mosquitoes in the field than other traps with
similar designs (Kline, 2002). The fans were powered by a
12 V battery and 100% CO2 was provided by a pressurized
cylinder releasing CO2 at 500 mL/min, which emulates the
output of the above MMF trap (Alan Grant, pers. comm.).
Air is pulled into the bottom of the trap, into a clear plastic
jar (about 11.4 L), through a screen and pushed out through
the top of the jar. Mosquitoes that are pulled in collect in the
jar. Because this trap has a counterflow design similar to the
MMF, except that it releases CO2 without heat or humidity,
initially it was chosen to examine the difference between the
presence and absence of these additional cues. Inserts con-
taining octenol can be used with all three of the ABC traps
and have been found to improve trap capture for some
species of mosquitoes (Kemme et al., 1993). However, we
wished to compare traps releasing only CO2. Moreover,
addition of octenol to CO2 generally does not improve trap
catch of ornithophagous species, including several Culex
species (Gibson & Torre, 1999). Furthermore, octenol
added to CO2-baited traps reduced capture of Culex pipiens
pipiens L. (Burkett et al., 2001) and Cx. quinquefasciatus
(Russell, 2004).
To facilitate three-dimensional (3-D) video recording,
pressurized tanks were placed outside of the wind tunnel
for the MMF and MMX traps and against the inside wall
of the wind tunnel for the MML trap. Analysis of the
plume structures of CO2 and discussion of other physical
differences between these four traps can be found in
Cooperband & Cardé (2006).
Insects
A colony of anautogenous Cx. tarsalis was maintained in
Boyden Laboratory at University of California Riverside at
LD 16 : 8 h, 25 C and about 65% relative humidity. Eggs
were collected and placed in plastic water pans, and alfalfa
pellets for rabbits were added to the water as larval diet.
Fourth instar larvae of Cx. quinquefasciatus were obtained
weekly from the Walton Laboratory at University of
California Riverside, where they were reared at LD
16 : 8 h, at about 26 C, with 50% relative humidity (see
Georghiou & Wirth, 1997) and a larval diet consisting of
one part brewer’s yeast to three parts ground mouse chow.
Pupae of both species were collected daily and allowed to
emerge in screened cages (30
30
30 cm) with access to
25% honey solution and water. Colonies were maintained
by weekly blood feeding on live chicks, but mosquitoes
used in experiments had no prior blood-feeding experience.
In preliminary experiments it was determined that only 2%
# 2006 The Authors
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
Orientation of Culex mosquitoes to traps 13
and 9% of 3-day-old Cx. tarsalis and Cx. quinquefasciatus
females, respectively, flew upwind to EVS traps in the wind
tunnel, whereas 75% and 93% of 7–10-day-old females flew
upwind. Therefore, anautogenous female Cx. tarsalis or
Cx. quinquefasciatus, about 10 days post-eclosion, were
used. To allow mating, females were housed with males
from eclosion until experiments.
Experimental set-up
About 8–10 h prior to testing, mosquitoes were deprived
of honey solution and water. About 1–2 h prior to experi-
ments, females were aspirated into release cages made of
PlexiglasÒ cylinders (5 cm
7 cm diameter) with mesh on
one end and a slit near the open end into which a paper
cover was inserted. Because most host-seeking behaviour
by these species occurs soon after dusk (Meyer et al., 1985,
1986), they were allowed to acclimatize in their release
cages outside in the shade at dusk, during which time the
wind tunnel was constructed for that evening and recording
equipment was set up. Mosquito releases began following
sunset and experiments were conducted over the next
1–2 h. Only one species and one trap type were tested
per night.
A covered release cage containing four to six mosquitoes
was placed in a 1 cm-wide rubber band stapled to the end
of a 1.2 m-long wooden rod (1.5 cm diameter). It was then
extended just inside the door of the wind tunnel with the
mesh screen facing into the wind and held there for about
60 s to allow mosquitoes to rest on the mesh. The paper on
the downwind end of the release cage then was removed
gradually and the cage was slowly extended to the middle of
the wind tunnel, keeping the same orientation. The cage
was held for about 10 s in the centre of the CO2 plume and
then slowly turned until the open end rotated to face
upwind, at which point the mosquitoes flew out. The cage
was held in place until the mosquitoes completed their
flights upwind (to avoid making any movements that
might interfere with their behaviour), after which it was
removed and a new cage was placed in the rubber band.
Recording flights
Each trap was set up in the centre of the upwind end of
the wind tunnel, approximately 30 cm from the upwind
screen, and mosquitoes were released 55 cm above the
ground and 270 cm downwind of the traps. The mosquitoes
were video-recorded using Sony Hi8 (EVO-550H) recorders
and two Sanyo black and white video cameras (VCB-
3512T), with a shutter speed of 1/60 s and 6-mm lenses
that were aimed from two different angles at mosquitoes
as they flew upwind toward the traps. The two cameras
were synchronized using an Event & Video Control Unit
(Peak Performance Technologies Inc., Englewood, CO,
U.S.A.). One video camera was always aimed from the
side of the tunnel toward the trap, showing the XZ plane.
14 M. F. Cooperband and R. T. Carde´
The other video camera was aimed about 90 from the first,
and because the height of the trap being recorded varied,
the second camera was positioned either on the ceiling of
the wind tunnel (XY), in a hole in the ground below the
trap (XY) or downwind near the release position (YZ), to
obtain a clear view of mosquitoes approaching the traps.
The two video cameras were aimed at the trap and the area
directly downwind, capturing about the last 150 cm of the
approach to the trap. Three Tracksys Ltd. (Nottingham,
U.K.) Infrared LED Illuminator light sources (arrays of 90
LEDs with 880 nm peak output) were aimed upwind at the
trap from behind the release point to illuminate the mos-
quitoes. A calibration object with 20 fixed 3-D coordinates
was placed in the area just downwind of the trap entrance
and video recorded prior to releasing the first mosquito.
The coordinates provided by this object were later digitized
and these allowed the software to calculate accurate 3-D
coordinates of mosquitoes flying through the same space.
Environmental conditions
Experiments were conducted at the edge of a wooded
area of a University of California Riverside experiment
station between May and November of 2000, 2001 and
2002. The average temperature at the start of data
collection was 24.6 C, with a range between 20.6 and
28.1 C. The average relative humidity was 63.9%, ranging
from 22 to 87%. The average barometric pressure was
760.2 mmHg, ranging between 757.4 and 763.5 mmHg.
Track discrimination
Not all mosquito flights observed on the video monitor
satisfied requirements for analysis. To be considered, the
flight had to start downwind and end upwind, and be
largely visible on both video records. Upwind flights were
defined as those that started approximately 60 cm or more
downwind of the trap and progressed to within 30 cm of
the trap or closer. Tracks in which the mosquito flew from
the upwind end of the tunnel to the downwind end were
excluded. Similarly, tracks in which the mosquito reached
the trap but the approach did not start far enough down-
wind were discarded.
Of the tracks meeting the above requirements, not all
were recorded with sufficient clarity for analysis. Because
two cameras were used from different angles, on occasion a
mosquito was visible in one camera, but behind the trap or
out of view in the other camera. Because no.3-D coordi-
nates could be calculated for those frames, this would
appear as a gap when the final 3-D track was calculated
by the computer program MOTUS (Peak Performance
Technologies, Inc., Centennial, CO, U.S.A.). If more than
30% of all frames of a track were gaps, that track was
excluded from further analysis. In the remaining tracks,
the ‘interpolate gaps’ function of MOTUS was used to cor-
rect for missing coordinates. Following this procedure, the
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
tracks were drawn by MOTUS and visually inspected and
compared to the tracks prior to interpolation. If a large
section of adjacent frames was missing in a given track, it
sometimes resulted in an evident distortion of the flight
path by the interpolation procedure, in which case such a
track was also removed from the analysis.
Analysis of capture rates
Each night of testing was considered a replicate for a
given trap and species. Three sets of numbers for statistical
comparisons for each trap and species were calculated:
number of flights per number released, number caught
(þ0.5) per number released and number caught (þ0.5) per
number of flights. These ratios were square root arcsin
transformed. A few instances occurred where the ratio of
flights to number of mosquitoes released was greater than
1, in which case they were adjusted to 1 to permit the
transformation. These means were compared using
two-tailed t-tests with unequal variance (Microsoft Excel,
2002 SP3).
Three-dimensional analysis of flight tracks
For each track, the resulting pairs of two-dimensonal
(2-D) flight tracks from two views were entered as digital
coordinates into MOTUS, which was then used to calculate
the 3-D flight path taken by each mosquito. The 3-D flight
coordinates were entered into an in-house computer pro-
gram (TRACK 3-D, version 2.2.6, written by Josep Bau),
which calculated a variety of flight parameters in one, two
or three dimensions, such as: X speed (along wind velocity),
Y speed (across wind velocity), Z speed (vertical velocity),
track angle (XY), course angle (XY and 3-D), drift angle
(XY and 3-D), inclination angle (XZ), front angle (YZ),
ground speed (XY), airspeed (XY and 3-D), time elapsed,
flight angle (3-D), flight speed (3-D), vertical position (Z)
and tortuosity (XY, XZ, YZ and 3-D). Figures 1 and 2
provide diagrams of some 2-D and 3-D flight parameters.
Track angle (XY), inclination angle (XZ) and front angle
(YZ) represent the angle of movement of the mosquito in
the three respective planes. The flight angle is the angle of
movement of the mosquito in three dimensions (calculated
Fig. 1. The triangle of velocities as defined by Marsh et al. (1978)
looking down from above a mosquito (in the XY plane).
# 2006 The Authors
 Orientation of Culex mosquitoes to traps 15

Flights tracks were divided into 10-cm increment bins

Fig. 2. Definitions of angles and vectors used to calculate three-
dimensional flight parameters with the mosquito moving from the
box at point 0 to point 1 (flight vector). Shadows indicate the
mosquito’s location in the three planes.
using the flight vector). Drift angle is the angle of displace-
ment of the mosquito between its heading and its actual
trajectory as a result of the wind speed and direction.
Tortuosity is a measure of path straightness and, when
an organism is orienting toward a ‘goal’, indicates the
level of efficiency of the orientation mechanism being
used (Benhamou, 2004). In this study, tortuosity was cal-
culated by dividing the distance travelled between the
starting point and the end point by the straight distance
between those two points. The tortuosity index, therefore,
is always  1, and the higher the value, the more turning
occurred.
along the X-axis, starting at the downwind edge of the
trap, and flight parameters for each track were calculated
for each bin. For each species, flight parameters were com-
pared by trap. Flight parameters were also examined for
each species and trap to look for differences between mos-
quitoes that were caught or not caught.
Track durations for flight tracks of each trap and species
were compared by date using the Games–Howell post hoc
analysis of variance with P ¼ 0.05 (Games & Howell,
1976). This test was selected because it does not assume
equal sample sizes or equal variances. No significant differ-
ences were found between dates for each trap and species,
and so dates within each trap and species were combined
and individual tracks were used as replicates. Means of
flight parameters per bin were quantified for individual
mosquitoes, grouped by species and by trap, and compared
and tested for significance using the Games–Howell
post hoc analysis of variance with a P-value of 0.05.
Additionally, flight tracks of Cx. quinquefasciatus that
were caught were compared to flight tracks of those that
were not caught for each trap and bin using the Games–
Howell post hoc analysis of variance with a P-value of
0.05. Complete track durations of Cx. quinquefasciatus
mosquitoes that were caught were compared to those
that were not caught using a Mann–Whitney test
(P ¼ 0.05) for each trap.
Results
Trap capture
Table 1 provides numbers of mosquitoes released and
number of release nights per trap and species (nine
release nights were discarded due to technical problems).

Table 1. Proportions of female (a) Culex quinquefasciatus and (b) Culex tarsalis released, flying upwind and captured for four CO2-baited
traps. Numbers in columns followed by the same letters are not significantly different from each other using two-tailed t-tests with unequal
variance (P < 0.05).
(a) Culex quinquefasciatus

Flights/releases Captures/flights Captures/releases n mosquitoes released N release nights

EVS 0.47a 0.13a 0.04a 320 4

MMF 0.44a 0.16a 0.05a 423 6
MML 0.67a 0.14a 0.09a 330 5
MMX 0.54a 0.58b 0.26b 377 4

(b) Culex tarsalis

Flights/releases Captures/flights Captures/releases n mosquitoes released N release nights

EVS 0.75a 0.08a 0.06a 79 4

MMF 0.48a 0.37a 0.18a 152 2
MML – – – – –
MMX 0.24 0.71 0.17 58 1

EVS, Encephalitis Virus Surveillance trap; MMF, Mosquito Magnet Freedom trap; MML, Mosquito Magnet Liberty trap; MMX, Mosquito
Magnet-X trap.

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Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
16 M. F. Cooperband and R. T. Carde´

Table 2. The number of tracks of each treatment used for the with this species; the MMX trap was only tested on one
TRACK 3-D analysis of flight parameters. evening, and so it was not included in the trap-capture
comparison.
Trap

Species Caught? EVS MMF MML MMX

Number of tracks used
Culex quinquefasciatus No 72 42 61 50
Yes 3 3 7 11 Of 924 flights recorded (some mosquitoes returned to
Culex tarsalis No 42 19 – 7 the release area and flew upwind again), 536 satisfied
Yes – 4 – 3 the digitizing criteria (where the mosquito started and
finished and was visible in both camera views), and of
EVS, Encephalitis Virus Surveillance trap; MMF, Mosquito
those digitized, 324 were of sufficient clarity (< 30%
Magnet Freedom trap; MML, Mosquito Magnet Liberty trap;
MMX, Mosquito Magnet-X trap.
gaps and tracks not distorted) to analyse for flight para-
meters. The number of tracks used in each treatment for
3-D flight track analysis is presented in Table 2. None
For each species and trap, the proportion of female of the flight tracks of Cx. tarsalis resulting in capture by
mosquitoes that flew upwind and were caught are also the EVS trap satisfied the criteria for 3-D analysis.
indicated. Although approximately 50% of the mosqui- Because tracks differed in length and each started and
toes released flew upwind toward the four trap types, ended at different distances from the trap, each bin
only a small fraction of mosquitoes that were released, contained a different number of tracks. The numbers
or of those that flew upwind, were captured. The MMX of flight tracks per bin for each species and trap are
trap had the highest capture rate. For given in Fig. 3.
Cx. quinquefasciatus, 13–16% of upwind flights resulted
in capture for the EVS, MMF and MML traps, whereas
58% of flights to the MMX trap resulted in capture Track duration
(Table 1). For Cx. tarsalis, the percentage of upwind
flights resulting in capture did not differ between the On average, both Cx. quinquefasciatus and Cx. tarsalis
EVS and MMF traps. The MML trap was not tested spent more time orienting to the EVS trap than to

Fig. 3. Number (N) of mosquito flight

tracks per bin (distance downwind of trap
over 10-cm increments) for (a) Culex quin-
quefasciatus and (b) Culex tarsalis used in
analysis of flight parameters for different
trap types. EVS, Encephalitis Virus
Surveillance; MMF, Mosquito Magnet
Freedom; MML, Mosquito Magnet
Liberty; MMX, Mosquito Magnet-X.

# 2006 The Authors

Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
 Orientation of Culex mosquitoes to traps 17

Average Durations of
Flights To Different Traps
30
a

Fig. 4. Average number of seconds (þ SE) a

mosquitoes spent in upwind flights to differ- 20

Time (s)
ent traps for Culex quinquefasciatus and
Culex tarsalis. Numbers under bars are the
number of tracks analysed for each trap and b
10 b b
species. Different letters above bars represent
b
significant differences between traps for that
species using the Games–Howell test c
(P < 0.05) (N.T. ¼ not tested). EVS,
Encephalitis Virus Surveillance; MMF, 0
Mosquito Magnet Freedom; MML, N= 74 45 68 61 42 23 N.T. 10
Mosquito Magnet Liberty; MMX, EVS MMF MML MMX EVS MMF MML MMX
Mosquito Magnet-X. Cx. quinquefasciatus Cx. tarsalis

the MMF, MML and MMX traps (Fig. 4). For MMF
and MML traps, tracks of Cx. quinquefasciatus females
that were caught were significantly longer in duration
than females that were not caught (Fig. 5). Similar
trends were noted for EVS and MMX traps
(Cooperband, 2005).
Significance of parameters
Tracks of mosquitoes flying to different traps may or
may not differ significantly, depending on the parameter
considered. Additionally, significant differences detected
between traps may be detected at one distance downwind
of the trap but not at another. For each bin downwind
of the traps, the parameters for which significant differ-
ences were found between traps are presented in
Tables 3 and 4. Orientation behaviour did not appear
to differ between traps at distances greater than about
100 cm downwind of the traps. For Cx. quinquefasciatus,
the number of significantly different parameters was
greatest between the trap and 30 cm downwind of the
trap, and between 80 and 90 cm downwind of the traps.
For Cx. tarsalis, the number of significantly different
parameters was greatest between 20 and 30 cm down-
wind of the traps.
As the mosquito approached, some parameters had
more significantly different bins between traps than
others. Likewise, the number of significantly different
parameters between traps in each bin varied as well.
For both species, parameters with the most bins con-
taining significant trap differences were vertical position
and speeds involving the X-axis (flight speed, ground
speed, X speed, airspeed and 3-D airspeed). Fewer sig-
nificant differences were seen in parameters such as time
elapsed and angles associated with the XY plane (drift
angle, 3-D drift angle, track angle, flight angle and 3-D
course angle). The fewest differences were found in
parameters involving the Z axis (Z speed, inclination
angle and front angle) and tortuosity. Diagrams of para-
meter differences not shown here can be found in
Cooperband (2005).

Average Duration of Cx. quinquefasciatus

Complete Flight Tracks to Traps
60
Caught
Not caught
Track Duration (s)

40 *
Fig. 5. Track durations for Culex quinque-
fasciatus mosquitoes that were caught com-
pared to those that were not caught for
individual traps. Significant differences 20 *
are indicated with asterisks using
Mann–Whitney test (P < 0.05). EVS,
Encephalitis Virus Surveillance; MMF,
Mosquito Magnet Freedom; MML,
Mosquito Magnet Liberty; MMX, 0
Mosquito Magnet-X. EVS MMF MML MMX

# 2006 The Authors

Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
18 M. F. Cooperband and R. T. Carde´

Table 3. Significant differences for various flight parameters as female Culex quinquefasciatus mosquitoes orientated to different traps. Traps
are represented by the letters e (EVS), f (MMF), l (MML) and x (MMX). Data in columns are aligned between column headings, indicating
where the bin measurements were taken. Dashes separate significantly different traps for the particular flight parameter and distance
downwind of the trap using the Games–Howell test (P < 0.05), with significantly lower values on the left-hand side and higher values on
the right-hand side of each dash. When no significant differences were detected between any of the traps, the space was left blank. Distances
relate to the downwind edge of the trap entrance, and parameters for that column were calculated for each 10-cm increment downwind from
that point.
Culex quinquefasciatus

Distance downwind of trap (cm)

Parameter 150 140 130 120 110 100 90 80 70 60 50 40 30 20 10 0 10 20

Flight speed (3-D) x-l x-ef f-lx ef-l f-lx ef-l; f-x ef-lx ef-lx e-flx ef-x e-flx e-lx
Ground speed (XY) f-lx; e-l ef-l ef-l ef-l ef-lx ef-lx e-flx e-fx; f-x e-flx e-lx
X speed f-l f-lx ef-l f-l ef-l ef-lx ef-lx e-flx e-flx; f-x e-flx
Airspeed (XY) f-l f-lx ef-l f-l ef-l ef-lx ef-lx e-flx e-flx; f-x e-flx
3-D airspeed f-l f-lx ef-l f-l ef-l ef-lx ef-lx e-flx e-flx; f-x e-flx

3-D drift angle l-x l-f x-e flx-e flx-e flx-e flx-e
Drift angle (XY) l-x l-f x-e flx-e flx-e flx-e flx-e
Time elapsed f-e x-e fx-e fx-e lx-e
Track angle (XY) e-l x-l ex-l x-l
Flight angle (3-D) e-lf e-l x-l
3-D course angle e-l x-f x-l
Course angle (XY) x-l
Y speed x-f x-e x-f

Vertical position l-ex; f-x l-ex l-x el-x l-f l-efx

Z speed f-e e-lx e-lx
Inclination angle (XZ) f-e x-e e-x
Front angle (YZ) f-el e-x

3-D tortuosity flx-e flx-e

XY tortuosity flx-e flx-e
XZ tortuosity flx-e flx-e
YZ tortuosity fl-e

EVS, Encephalitis Virus Surveillance trap; MMF, Mosquito Magnet Freedom trap; MML, Mosquito Magnet Liberty trap; MMX, Mosquito
Magnet-X trap.

Velocity
Mosquitoes reduced their speed as they approached
traps. Several patterns were apparent when considering
the number of parameters with significant differences at
various distances downwind of the traps. At the farthest
downwind distances, few parameters differed between traps
for either species (Tables 3 and 4). Starting at 90 or 100 cm
downwind of the traps, Cx. quinquefasciatus flew signifi-
cantly faster to the MML trap than the MMF trap, and
this continued until 20–30 cm downwind, where they both
slowed to similar velocities. Flight speed, ground speed,
3-D airspeed and X speed were similar (Cooperband,
2005), and so only X speed is presented for
Cx. quinquefasciatus and Cx. tarsalis (Fig. 6). As mosqui-
toes flew from 50 to 30 cm downwind of traps, they flew
significantly slower along the X-axis to the EVS and MMF
traps than to the MML or MMX traps, as indicated for all
speed parameters involving the X-axis (Tables 3 and 4). Y
speed and Z speed have few significant differences at dif-
ferent distances from traps (Tables 3 and 4). For all four
traps, mosquitoes did not vary their speed along the Y-axis,
but speed along the Z-axis increased from close to zero to
about 10 cm/s, within 50 cm downwind of the traps, indi-
cating that they flew upward more rapidly when close to the
traps (Cooperband, 2005). As mosquitoes approached the
traps, velocity parameters involving the X-axis show that
both species flew more slowly to the EVS trap than to any
of the others. This continued all the way to the leading edge
of the traps (0 cm downwind of trap) for both species.
There were no major patterns of differences between the
traps for Cx. tarsalis upwind of the leading edge of the
traps, but Cx. quinquefasciatus upwind of the EVS trap
continued to have a significantly lower velocity than with
the MML or MMX traps.
Angular headings
Drift angle (XY) and 3-D drift angle seemed to reveal
more differences between traps than track angle (XY),
inclination angle (XZ) and front angle (YZ) for both

# 2006 The Authors

Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
 Orientation of Culex mosquitoes to traps 19

Table 4. Significant differences for various flight parameters as female Culex tarsalis mosquitoes orientated to different traps. Traps are
represented by the letters e (EVS), f (MMF), l (MML) and x (MMX). Data in columns are aligned between column headings, indicating where
the bin measurements were taken. Dashes separate significantly different traps for the particular flight parameter and distance downwind of
the trap using the Games–Howell test (P < 0.05), with significantly lower values on the left-hand side and higher values on the right-hand side
of each dash. When no significant differences were detected between any of the traps, the space was left blank. Distances relate to the
downwind edge of the trap entrance, and parameters for that column were calculated for each 10-cm increment downwind from that point
Culex tarsalis.

Distance downwind of trap (cm)

Parameter 150 140 130 120 110 100 90 80 70 60 50 40 30 20 10 0 10 20

Flight speed (3-D) e-fx ef-x e-x ef-x ef-x ef-x ef-x ef-x
Ground speed (XY) e-fx ef-x e-x ef-x ef-x ef-x e-x ef-x
X speed e-fx ef-x e-x ef-x ef-x ef-x ef-x e-x
Airspeed (XY) e-fx ef-x e-x ef-x ef-x ef-x ef-x e-x
3-D airspeed e-fx ef-x e-x ef-x ef-x ef-x ef-x e-x

3-D drift angle x-fe x-fe x-fe x-fe x-fe x-fe x-e xf-e
Drift angle (XY) x-fe x-fe x-fe x-fe x-e x-e
Time elapsed f-e x-e xf-e; x-f x-fe xf-e xf-e
Track angle (XY) e-f
Flight angle (3-D) e-f
3-D course angle e-f
Course angle (XY)
Y speed

Vertical position f-e f-e f-e f-e f-e fx-e fx-e fx-e fx-e x-e fx-e fx-e
Z speed
Inclination angle (XZ)
Front angle (YZ)

3-D tortuosity x-e

XY tortuosity
XZ tortuosity
YZ Tortuosity xf-e

EVS, Encephalitis Virus Surveillance trap; MMF, Mosquito Magnet Freedom trap; MML, Mosquito Magnet Liberty trap; MMX, Mosquito
Magnet-X trap.

species (Tables 3 and 4). For both species, the MMX drift
angles are significantly smaller than the EVS drift angles,
whereas the course angles are not. The difference in signifi-
cance between drift angles and course angles can be
explained by the fact that during upwind flight, without
altering ground speed and wind speed, as the track angle
approaches 90 , the drift angle will change much more than
the course angle.
Vertical position
For both species, although other flight parameters invol-
ving the Z-axis produced few significant differences
between traps, the vertical position of mosquitoes as they
flew upwind toward the traps produced a relatively large
number of significant differences between traps (Tables 3
and 4; Fig. 7). Cx. quinquefasciatus females approached all
four traps from below the trap entrances. Although
Cx. tarsalis females followed the same approach pattern
for the MMF and MMX traps, as they neared the EVS
trap, they approached closer to the CO2 outlet.
Tortuosity
Although tortuosity differed significantly in only a few
bins, those bins are probably the most relevant in terms of
insects being trapped, because the mosquitoes are then
close to the trap entrance. Calculations of 3-D tortuosity
can be seen for both species in Fig. 8, and significant dif-
ferences between traps are noted in Tables 3 and 4. Both
mosquito species have non-tortuous flights upwind until
they arrive at the trap entrances. Cx. quinquefasciatus
females have significantly more tortuous flights 10–30 cm
downwind of the EVS trap than any other trap. Cx. tarsalis
females follow a similar pattern, but the differences are not
significant, probably due to the large variation and the
small N.
Time elapsed
From about 50 cm downwind to the leading edge of the
traps, Cx. quinquefasciatus females spent significantly more
time orienting toward the EVS trap than the MMF, MML

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Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
20 M. F. Cooperband and R. T. Carde´

Fig. 6. Average velocity along the X-axis of

(a) Culex quinquefasciatus and (b) Culex
tarsalis females at distances of 10-cm incre-
ments downwind of four types of mosquito
traps. EVS, Encephalitis Virus Surveillance;
MMF, Mosquito Magnet Freedom; MML,
Mosquito Magnet Liberty; MMX,
Mosquito Magnet-X.

Fig. 7. Average vertical positions of (a)

Culex quinquefasciatus and (b) Culex tarsa-
lis females with respect to the trap entrance
(0,0), at distances of 10-cm increments
downwind of four types of mosquito traps.
EVS, Encephalitis Virus Surveillance;
MMF, Mosquito Magnet Freedom; MML,
Mosquito Magnet Liberty; MMX,
Mosquito Magnet-X.

# 2006 The Authors

Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26

Fig. 8. Average three-dimensional (3-D)
tortuosity of (a) Culex quinquefasciatus
and (b) Culex tarsalis females at distances
of 10-cm increments downwind of four
types of mosquito traps. EVS, Encephalitis
Virus Surveillance; MMF, Mosquito
Magnet Freedom; MML, Mosquito
Magnet Liberty; MMX, Mosquito
Magnet-X.
or MMX traps (Fig. 9a, Table 3), and Cx. tarsalis females
similarly spent significantly more time orienting toward the
EVS or MMF traps than the MMX trap (Figs 4 and 9b,
Table 4). For both species and all traps, more time was
spent in bins near the trap entrances than further
downwind.
Differences between caught and not caught
For Cx. quinquefasciatus, flight tracks of mosquitoes
that were caught were compared to those that were not
caught for each trap. Table 5 shows which traps had
differences between caught and not caught mosquitoes
for each flight parameter and distance downwind of the
trap. Again, parameters involving speed along the X-axis
as well as vertical position often showed significant
differences followed by angular headings involving the
Y-axis, then other parameters involving the Z-axis, and
finally tortuosity. Mosquitoes that were caught had
significantly lower velocities and greater drift angles
than those that were not caught. Diagrams comparing
values for some of these flight parameters between
caught and not caught mosquitoes can be found in
Cooperband (2005).
Table 5 shows that significant differences between
flights for mosquitoes that were caught or not caught
occurred in different bins for different traps. The EVS
trap flights differed between 20 and 50 cm downwind of
the trap, the MMF trap flights differed between 0 and
20 cm downwind of the trap, the MMX trap flights
# 2006 The Authors
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
Orientation of Culex mosquitoes to traps 21
differed from 10 cm downwind of the trap to 20 cm
upwind of the trap, and the MML flights did not differ
in as clear a pattern, but several differences occurred
between 40 and 50 cm downwind of the trap. There
were few differences between Cx. quinquefasciatus
females that were caught and not caught in terms of
the height relative to the trap entrance, except for those
flying to the MML trap, in which the mosquitoes that
were caught approached the trap from farther below the
trap entrance than those that were not caught. For the
MMF trap the XZ tortuosity 0–20 cm downwind of the
trap was greater for mosquitoes that were caught than
those that were not caught.
Discussion
This is the first study to document the proportion of mos-
quitoes approaching CO2-baited traps that is captured and
to describe their orientation behaviour during their
approach. Some traps were surprisingly inefficient, captur-
ing less than 10% of the Cx. quinquefasciatus and up to
18% of Cx. tarsalis mosquitoes released. The MMX trap
was the most efficient for Cx. quinquefasciatus, with cap-
ture of 26% of the mosquitoes released. Additionally,
13–16% of Cx. quinquefasciatus females that flew upwind
to EVS, MMF and MML traps, and 58% of those
flying upwind to the MMX trap, were captured, and the
results for Cx. tarsalis were similar. Similarly, in a field
comparison, the MMF captured fewer mosquitoes than
the counterflow geometry trap (MMX), although both
22 M. F. Cooperband and R. T. Carde´
captured more than a CDC-style trap similar to the EVS
trap used in this study (Burkett et al., 2001).
Flight tracks of both species to the EVS trap were sig-
nificantly longer in duration than any of the other traps,
suggesting that the EVS trap attracts mosquitoes, but not
necessarily to the vicinity where they would be captured.
Furthermore, Cx. quinquefasciatus mosquitoes that were
caught spent more time orienting to traps than those that
were not caught, suggesting that whether a mosquito is
captured depends on how much time it is willing to invest
flying around the trap before giving up (Figs 4 and 5).
Although the MMX trap releases neither heat nor
humidity, it still outperformed the EVS, MMF and MML
traps with the two species tested. As shown in Cooperband
& Cardé (2006), the concentration of CO2 produced by the
MMX was intermediate between that of the EVS trap and
the other ABC traps, and its plume structure resembles that
of the MMF trap. The greatest incidents of bursts (when
CO2 concentration rises above background levels) for the
three ABC traps generally occurred below the height of the
trap entrances, whereas for the EVS trap burst numbers
were greatest at the level of the trap entrance and above,
especially at distances closer to the trap. The airspeed at the
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
Fig. 9. Average time elapsed for (a) Culex
quinquefasciatus and (b) Culex tarsalis
females at distances of 10-cm increments
downwind of four types of mosquito traps.
EVS, Encephalitis Virus Surveillance;
MMF, Mosquito Magnet Freedom; MML,
Mosquito Magnet Liberty; MMX,
Mosquito Magnet-X.
suction inlet and CO2 outlet was higher for the MMX trap
than any of the other traps, which probably contributed to
its higher trapping efficiency. The entrance of the EVS trap
was 20 cm away from the CO2 source, as opposed to 10 cm
for the ABC traps, but this did not seem to significantly
reduce its efficiency compared to the MMF and MML
traps, possibly because other factors such as visual cues
may have reduced trap capture for those traps.
For all trap types and both Culex species, flights were
relatively straight and rapid at distances greater than
100 cm downwind of the trap. Within 100 cm of the trap,
different mosquito orientation behaviour was observed
with each trap type. Mosquitoes approaching the EVS
and MMF traps tended to increase the tortuosity of their
flight, and spent more time within 30 cm of reaching the
trap. Mosquitoes approaching the MMX trap flew rela-
tively straight until passing the leading edge of the trap, at
which point they engaged in more tortuous flight.
Figures in Cooperband (2005) show the angular headings
for both species, how the 3-D drift angles of Cx. quinque-
fasciatus females are greater when flying to the EVS trap
than the other traps from 40 cm downwind to the leading
edge of the traps, and how for Cx. tarsalis females
# 2006 The Authors
 Orientation of Culex mosquitoes to traps 23

Table 5. Significant differences between caught and not caught Culex quinquefasciatus females for various flight parameters at different
10-cm increments downwind of four different traps. Traps are represented by the letters e (EVS), f (MMF), l (MML) and x (MMX). Data in
columns are aligned between column headings, indicating where the bin measurements were taken. A letter indicates a significant difference
between caught and not caught tracks for that trap, and for the particular flight parameter and distance downwind of the trap, using the
Games–Howell test (P < 0.05). When no significant differences were detected, the space was left blank. Distances relate to the downwind
edge of the trap entrance.
Culex quinquefasciatus – differences between those caught and not caught

Distance downwind of trap (cm)

Parameter 100 90 80 70 60 50 40 30 20 10 0 10 20

Flight speed (3-D) l x fx

Ground speed (XY) l x e fl fx x
X speed e e e f fx x
Airspeed e e e f fx
3-D airspeed e e e f fx

3-D drift angle fx x x

Drift angle (XY) l e f x x
Time elapsed l x x
Track angle l e e
Flight angle l e e
3-D course angle l e
Course angle l e e l
Y speed l

Vertical position l l l l l l
Z speed x f
Inclination angle ef
Front angle l

3-D tortuosity l
XY tortuosity
XZ tortuosity l f
YZ tortuosity x

EVS, Encephalitis Virus Surveillance trap; MMF, Mosquito Magnet Freedom trap; MML, Mosquito Magnet Liberty trap; MMX, Mosquito
Magnet-X trap.

orienting to the MMX trap 3-D drift angle is lower than to
the other traps from 70 cm downwind to the leading edge
of the trap.
Mosquitoes approaching the MML trap appear to have
very little tortuosity. The MML trap presented the most
logistical problems in terms of recording 3-D video near
the trap entrance. This lowered the N near that entrance
and could be partly responsible for the low tortuosity read-
ings near the entrance of that trap. For all traps, few bins
were significantly different for tortuosity, but this may be
explained by the fact that generally the mosquitoes flew
fairly straight tracks until they reached the trap, at which
point they spent more time meandering around before being
caught or leaving (see example track in Fig. 10). This
appeared to be the case for all the traps except for the
MMX. Mosquitoes seemed to fly directly to the entrance
of the MMX and become captured, or fly past it and turn
around. As some species of mosquitoes reach the source of
CO2, they may shift orientation strategies and fly to promi-
nent visual cues during their final approach (Gibson & Torr,
1999), and the fact that the MMX trap was mostly transpar-
ent may have played a role in its relative success. This is most
evident when looking at the time elapsed and tortuosities for
Cx. quinquefasciatus. Mosquitoes spent more time slowly
flying around and turning immediately downwind of the
entrances of the EVS, MMF and MML traps, whereas this
behaviour was rarely observed near the MMX trap. In fact,
there is a marked increase in tortuosity after the mosquitoes
passed the leading upwind edge of the MMX trap that is
absent in approaches to the other three traps. This suggests
that mosquitoes flying to the MMX trap may not perceive
some cues that would cause them to reduce flight velocity,
but after passing the trap, they exit the CO2 plume, and then
switch to a ‘meandering’ behaviour that may improve their
chance of regaining contact with the plume. Mosquitoes that
continue to follow the plume, rather than perhaps flying to a
prominent visual cue, would be ‘led’ along the plume to its
source close to the trap entrance, increasing the likelihood of
capture. We did not see a significantly different prominent
peak in tortuosity downwind of traps for Cx. tarsalis. In
fact, there is a peak just upwind of the EVS trap for this
species. It is possible that this species relies more upon the
odour plume than on visual cues, even during the final
approach to the source.

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Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
24 M. F. Cooperband and R. T. Carde´
XZ
X
XY
Because Cx. tarsalis females approached all traps at the
height of the CO2 outlet, it appears that they may be relying
principally on the location of the CO2 plume, as opposed to
Cx. quinquefasciatus females, which approached all traps
from below the trap entrances (Fig. 7). Mosquitoes that
approached the MMF trap were sometimes seen meander-
ing slowly around its metal base, which may explain why
upwind of this trap the average altitudes of both species
were reduced considerably.
Although Cx. pipiens (sensu lato) is reportedly highly
ornithophagous (Bohart & Washino, 1978), the host
preference of Cx. quinquefasciatus is more anthropophilic.
Samuel et al. (2004) reported that 75% of Cx. quinquefas-
ciatus females collected indoors had fed upon humans, and
only about 2% had fed on birds. Of the combined
Cx. quinquefasciatus mosquitoes captured both indoors
and outdoors, Gomes et al. (2003) reported that about
52% and 22% had fed on humans and chickens, respec-
tively. By contrast, many studies on the host feeding pat-
terns of Cx. tarsalis show that this species feeds on a broad
range of hosts but prefers birds (Bohart & Washino, 1978).
In a field study, Wekesa et al. (1997) found that Cx. tarsalis
fed upon birds and mammals about 71% and 27% of the
time, respectively. If Cx. tarsalis females follow plumes to
their source, relying more on olfactory and less on visual
cues, this may aid in the location of birds as hosts, which
provide less conspicuous visual cues in the dark at close
range than humans. Similarly, if Cx. quinquefasciatus
females rely more on visual cues when close to their hosts,
this may aid in finding their preferred larger mammalian
hosts and reduce their success at finding birds.
Furthermore, the preference of biting sites around the
human torso and legs by Cx. quinquefasciatus (de Jong &
Knols, 1995) suggests that at some point during their
approach to a human host, they would depart from the
Journal compilation # 2006 The Royal Entomological Society, Medical and Veterinary Entomology, 20, 11–26
YZ
Fig. 10. A schematic representation of an
actual track of a Culex tarsalis female flying
upwind to an Encephalitis Virus
Surveillance (EVS) trap and not getting
caught, showing the same track in a three-
dimensional view and in the three different
planes.
plume of CO2 originating from the head, which is the least
preferred biting site for that species.
The traps examined had multiple variables and therefore
we can only consider which features correlate with particu-
lar orientation manoeuvres and we cannot verify cause and
effect. Although there are several differences between the
three ABC traps, a common feature was the CO2 outlet
being positioned 10 cm below the trap entrance, whereas on
the EVS trap it was located 20 cm above the trap entrance.
This factor seemed to play a more important role in the
orientation behaviour of Cx. tarsalis females, which
appeared to follow the location of the CO2 plume with the
highest burst numbers to the source which was 20 cm higher
than the entrance in the case of the EVS trap, while
Cx. quinquefasciatus females approached the trap on average
from below the height of the trap entrance for all four traps.
Our observations of approaching mosquitoes that were
and were not captured by the EVS trap, suggest that
capture might have been occurring randomly, in that a
captured mosquito engaged in a meandering behaviour
that could last several minutes just downwind of the trap,
eventually happened to get too close to the entrance.
Mosquitoes that ‘gave up’ sooner therefore tended not to
be captured. Alternatively, some mosquitoes flying to the
MMF, MML and especially the MMX trap sometimes flew
directly into the trap entrance, without a long bout of first
meandering back and forth.
Difference in maximum concentration of CO2 between
traps did not appear to affect orientation behaviour
(Cooperband & Cardé, 2006). If the prediction by Gillies
(1980) that the most important feature of the plume for
orienting mosquitoes may be changes in CO2 concentration
is correct, then burst number, which would be detected by the
mosquito as a change in concentration, would be a more
important feature of the plume than peak concentration. In
# 2006 The Authors

Cooperband & Cardé (2006) we found that, at three CO2
thresholds, burst number was low 60 cm downwind of the
EVS trap at the height below the trap entrance, whereas it
remained high, at least at one of the three thresholds, for the
ABC traps to 30 cm downwind of the traps at the height
below the trap entrances. This comparatively low burst num-
ber frequency 60 cm downwind of the EVS trap below the
trap entrance is likely caused by the CO2 outlet being farther
away from the trap entrance on the EVS trap than on the
ABC traps, and the fact that the CO2 is released passively
rather than being blown downward. The low burst frequency
may explain why mosquito flights downwind of the EVS trap
became much more tortuous, resulting in more time elapsed
prior to capture. Cooperband & Cardé (2006) found that the
plume of the MMX trap, unlike the MMF and MML plumes,
dropped below the downward-pointing trap entrance, which
may have contributed to the improved capture rate of that
trap. Another possible explanation for the low capture effi-
ciencies of these traps is that they may lack other important
cues such as additional semiochemicals. Further behavioural
studies are required in which some of the variables (such as
relative plume location, suction strength, burst number, or
visual appearance) can be varied systematically.
Important caveats for this study are first that these findings
apply only to the orientation of the two species studied,
Cx. quinquefasciatus and Cx. tarsalis, in a light and unvarying
wind. In relatively still air or in a turbulent flow, capture rates
could be elevated or suppressed. Second, field trapping effi-
ciencies could be higher than reported here, if mosquitoes that
were not initially captured later reoriented to traps. Third,
adding of other host odours to CO2 might change orientation
behaviour, for example increasing the giving up time, which in
turn might provide more opportunity for mosquitoes to
meander close to the trap entrance and be captured.
Acknowledgements
Dr Alan Grant of American Biophysics Corporation and
the Northwest Mosquito and Vector Control District gene-
rously provided the traps. We thank Dr Bill Walton for
providing mosquitoes, Terence Fung, Ross Whittaker,
Andrew Kuszynski, Brian Panama and Michelle Sanford
for their assistance in experiments and data entry,
Kris Tollerup and Dr Robert Beaver for statistical advice,
Dr Josep Bau for his work on the TRACK 3-D computer
program, Dr Kris Justus for training and advice, and
Dr Marieta Braks for critical review. Drs Scott Ritchie
and Alan Grant provided useful comments. We also
acknowledge support from the University of California
Systemwide Mosquito Research Program.
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Orientation of Culex mosquitoes to traps 25
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Accepted 18 December 2005
# 2006 The Authors