December, 2004
Journal of Vector Ecology
309
The relative attractiveness of carbon dioxide and octenol in CDC- and
EVS-type light traps for sampling the mosquitoes Aedes aegypti (L.),
Aedes polynesiensis Marks, and Culex quinquefasciatus Say
in Moorea, French Polynesia
Richard C. Russell
Department of Medical Entomology, University of Sydney and ICPMR, Westmead Hospital,
Westmead, NSW 2145, Australia
Received 24 March 2004; Accepted 18 April 2004
ABSTRACT: Two dominant day-biting pests and vector species on the island of Moorea in French Polynesia are
Aedes (Stegomyia) aegypti (L.) and Aedes (Stegomyia) polynesiensis Marks, major vectors of dengue viruses and
Wuchereria bancrofti, respectively. Their surveillance is hindered by a relative lack of attraction to light traps,
necessitating the undesirable use of human bait collections with the inherent risks of pathogen transmission. The
effectiveness of CDC- and EVS-type light traps baited with olfactory attractants was evaluated for these two Aedes
species and the nocturnal Culex (Culex) quinquefasciatus Say in three sites in urban and semi-rural environments
on Moorea in October/November 2003. Firstly, four CDC-type traps with light only, light with octenol, light with
carbon dioxide (dry ice), and light with octenol plus carbon dioxide were operated continuously over four days with
daily rotation to compensate for position effects. Secondly, two CDC- and two EVS-type traps with carbon dioxide
or carbon dioxide plus octenol were operated continuously over four days with similar rotation. Variation was found
in the numbers of the three species collected at the different sites, reflecting the relative availability of their preferred
larval habitats. With the CDC traps in the first trial, the addition of octenol to the light did not significantly increase
the collection of any species, the addition of carbon dioxide did significantly increase collection of all three species,
while the addition of octenol to the light plus carbon dioxide did not significantly increase the collections further. In
the second trial, there was no significant difference in the mean number of Ae. aegypti or Ae. polynesiensis collected
in either EVS or CDC traps when baited with carbon dioxide or with octenol added. For Cx. quinquefasciatus, the
supplementation with octenol made no significant difference with EVS traps but resulted in significantly reduced
collections in CDC traps. Overall, neither trap, however baited, provided large samples when compared with landing/
biting collections at human bait. Only two other species were collected, Culex (Culex) roseni Belkin and Aedes
(Aedimorphus) nocturnus (Theobald), the latter being a first record for the island of Moorea and for French
Polynesia. Journal of Vector Ecology 29 (2): 309-314. 2004.
Keyword Index: Aedes aegypti, Aedes polynesiensis, CDC and EVS traps with carbon dioxide and octenol, French
Polynesia.
INTRODUCTION
The two dominant day-biting pest species on the
island of Moorea in French Polynesia, Aedes (Stegomyia)
aegypti (L.) and Aedes (Stegomyia) polynesiensis Marks,
are the major local vectors of dengue viruses and
Wuchereria bancrofti, respectively, while Culex (Culex)
quinquefasciatus Say is a common night-biting pest but
is generally considered a much less important vector of
filariasis in the Pacific regions (Lee et al. 1987, 1989).
Surveillance of the urban and peri-urban populations
of these Aedes mosquitoes for their abundance and
infection rates is problematic. For Ae. aegypti, indices
of larval abundance in artificial domestic containers do
not adequately indicate adult abundance and the attendant
risks of transmission (Tun-Lin et al. 1996). For Ae.
polynesiensis, its appearance in artificial containers in
residential environments does not indicate the more
extensive colonization of natural containers such as treeholes, palm fronds, coconuts, and crab holes and grossly
underrepresents adult activity. Additionally, adult
population densities of Ae. aegypti can be difficult to
estimate as they can be relatively low compared with
other species of mosquito (Reiter and Gubler 1997),
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Journal of Vector Ecology
particularly species such as Ae. polynesiensis that can
have myriad larval habitats available (Lee et al. 1987) and
Cx. quinquefasciatus which can proliferate in high
nutrient wastewater in urban areas (Lee et al. 1989).
Development of accurate methods for adult
surveillance of these two Aedes has been hindered by
the relative lack of attraction of both to light traps. This
problem has resulted in the use of human bait collections
as the standard sampling procedure, a technique that is
labor-intensive and influenced by a range of
environmental, site, and collector factors, and is also
undesirable because it exposes collectors to the risk of
pathogen transmission (Reiter and Gubler 1997).
However, in many situations where there is a range
of vector mosquito species, crepuscular and nocturnal
as well as diurnal species, the facility to employ a single
attractive trap is highly desirable for mosquito
surveillance. Accordingly, the effectiveness for collecting
Ae. aegypti and Ae. polynesiensis, and the other
dominant domestic pest species Cx. quinquefasciatus,
of two commonly used miniature light traps baited with
the olfactory attractants carbon dioxide and octenol was
assessed at strategic locations in urban and semi-rural
environments on Moorea. Human bait landing/biting
collections were used as a comparative reference
technique.
MATERIALS AND METHODS
The investigations were conducted during October
and November 2003 on the northern side of Moorea
Island in three localities from Cooks Bay to Opunohu
Bay: Pao Pao, University of California Berkeley Gump
Station, and Vaihara. Prior to the trials, human bait
collections at Pao Pao indicated many Ae. aegypti and
few Ae. polynesiensis were active and larval collections
confirmed a number of Ae. aegypti habitats nearby. At
Gump Station, human bait collections indicated many
Ae. polynesiensis and few Ae. aegypti adults were active
and larval searches confirmed a few Ae. aegypti habitats
nearby. At Vaihara, human bait collections confirmed both
Ae. aegypti and Ae. polynesiensis were active and larval
collections confirmed some Ae. aegypti habitats nearby.
Potential larval habitats for Ae. polynesiensis (e.g. leaf
axils, tree holes, coconuts and husks, crab holes) were
numerous in all three areas. Collections with a “gravid
trap” (Reiter, 1987) indicated Cx. quinquefasciatus was
present at all three sites.
The relative effectiveness of CDC-type traps
(BioQuip Products Miniature Light Trap Model 2836)
was investigated by setting the traps in groups of four
in each of the three localities, with one trap unbaited (i.e.
light alone) and the other traps with either octenol in a
December, 2004
glass vial with cotton pipe-cleaner as wick, as per Van
Essen et al. (1994), carbon dioxide (dry ice), or carbon
dioxide plus octenol. The traps were placed in windsheltered positions, operated continuously for 24 h a
day for four d (with octenol continuously available and
dry ice provided twice a day), and rotated between the
four positions on a daily basis to compensate for position
effects.
The comparative effectiveness of the CDC-type (as
above) and the Encephalitis Virus Surveillance (EVS)type (BioQuip Products Heavy Duty EVS CO2 Trap
Model 2801A) traps with octenol and/or carbon dioxide
was investigated with four traps (two of each type) in
each of the three localities. One of each type was baited
with carbon dioxide (dry ice) alone and the other was
baited with carbon dioxide plus octenol. The collection
containers used for both traps were identical and were
the solid cups supplied by BioQuip for their CDC-type
trap. The traps were placed in wind-sheltered positions,
operated continuously for 24 h a day for four d (with
octenol continuously available and dry ice provided
twice a day), and rotated between the four positions on
a daily basis to compensate for position effects.
Daily collections could not be separated for
logistical reasons, but this was not considered to be a
critical consideration because the principal objective
was simply to compare the collection of each species
with the various traps and attractants at the three
different locations. The numbers of each species of the
three mosquitoes collected in the first trial by each CDCtype trap baited with light, octenol, and carbon dioxide
across the three locations were log (x+1) transformed
then compared using one-way analysis of variance
(ANOVA). The numbers collected by the variously baited
CDC and EVS traps in the second trial were treated
similarly. For both analyses, Tukey’s pairwise multiple
comparison test was used to compare means.
In parallel with the above trials, human bait landing/
biting collections were undertaken for 15 min in the two
h prior to sunset, in the vicinity of the traps, to provide
a comparison of techniques.
RESULTS
In the first trial, there was great variation in the
numbers of the three species collected by the CDC traps
within and between the three sites (Table 1), reflecting
the relative local availability of preferred larval habitats
and adult production as indicated by the prior local
human bait collections and larval surveys, as mentioned
above. Aedes aegypti was the most abundant Aedes
species at Pao Pao, while Ae. polynesiensis was
predominant at Gump Station and Vaihara. Analysis of
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Journal of Vector Ecology
311
Table 1. Numbers of Aedes aegypti, Aedes polynesiensis, and Culex quinquefasciatus collected in CDC-type light
traps, and also baited with carbon dioxide (dry ice) and/or octenol (1-octen-3-ol), over 24 h for four d, in three
locations between Cooks Bay and Opunohu Bay, Moorea, French Polynesia, October 2003.
Species
Location
light only
Ae. aegypti
Ae. polynesiensis
Cx. quinquefasciatus
Pao Pao
Gump Stn
Vaihara
Pao Pao
Gump Stn
Vaihara
Pao Pao
Gump Stn
Vaihara
0
0
0
0
1
0
4
1
2
these data showed that the addition of octenol to the light
attraction of the CDC trap did not significantly increase
the collections of any species. The addition of carbon
dioxide to the light did significantly increase collections
of each species at all three sites, but the supplementation
of carbon dioxide with octenol did not increase
collections further (Table 2).
In the second investigation, the same variation in
relative abundance of the Aedes species at the three
sites was demonstrated (Table 3). Analysis of these data
showed no significant difference in mean numbers of
Ae. aegypti and Ae. polynesiensis collected by either
CDC or EVS traps, however they were baited. For Cx.
quinquefasciatus, there was no significant difference in
numbers collected by either trap baited with carbon
dioxide alone. The addition of octenol to the carbon
dioxide bait resulted in significantly reduced collections
with CDC traps but this impact was not seen with the
EVS traps (Table 4).
In comparison, the landing/biting collections at
human bait produced relatively larger numbers of both
Aedes species (but not Cx. quinquefasciatus) within the
smaller collection period. For Ae. aegypti, there were 15
females at Pao Pao, 3 females at Gump, and 4 females at
Vaihara per 15 min and for Ae. polynesiensis there were 4
females at Pao Pao, 17 females at Gump, and 22 females
at Vaihara, per 15 min collecting time.
Two other species were recorded in the trap
collections at Vaihara (Table 3), Culex (Culex) roseni
Belkin and Aedes (Aedimorphus) nocturnus, (Theobald),
the latter being a first record for the island of Moorea
and for all of French Polynesia.
CDC trap bait
light plus
light plus
light plus
octenol
carbon dioxide octenol plus CO2
1
29
21
2
5
6
0
11
6
2
7
2
3
17
10
2
16
12
1
13
5
1
5
2
1
6
4
DISCUSSION
These investigations occurred during October and
November, in the latter half of the ‘so-called’ dry season,
but some rain fell patchily during each month and
increased in amount and frequency over the three mo
from September. Populations of both species increased
from September through November along with the
increasing rainfall and humidity (data not included).
The operation of the traps throughout the 24 h day
was designed to allow the collection of mosquitoes
displaying diurnal, crepuscular, and nocturnal activity.
Aedes aegypti in Malaysia displayed its greatest biting
activity in mid-morning and late afternoon (Macdonald
1956), similar to that found in Kenya (Teesdale 1955). In
Trinidad, Chadee (1988) found diurnal peaks at 06000700 and 1700-1800 h with the morning peak being slightly
greater, but in a later report (Chadee and Martinez 2000)
described a trimodal pattern of landing with consistent
peaks at 0700, 1100, and 1700 h that was similar to the
trimodal pattern reported from Tanzania by Corbet and
Smith (1974) and from Indonesia by Atmosoedjono et al.
(1972) which had earlier been dismissed as due to
sampling error or seasonal or geographic factors.
Additionally, Ae. aegypti is well known for extended
host-seeking activity and for taking multiple blood meals
in a single gonotrophic cycle (Macdonald 1956, Scott et
al. 1993).
Aedes polynesiensis was found generally to be a
daytime biter in Fiji (Rakai et al. 1974), while in American
Samoa and Independent Samoa, Jachowski (1954) and
Suzuki and Sone (1973), respectively, reported the species
to be diurnally active with a major peak of activity in late
312
Journal of Vector Ecology
December, 2004
Table 2. Mean number (± S.D.) of Aedes aegypti, Aedes polynesiensis, and Culex quinquefasciatus collected in
CDC-type light traps and also baited with carbon dioxide (dry ice) and/or octenol (1-octen-3-ol) at Moorea, French
Polynesia, October 2003.
LIGHT
LIGHT+OCT
LIGHT+CO2 LIGHT+OCT+CO2
Ae. aegypti
0.33±0.58 a1
1.00±1.00 a
15.00±12.49 b
11.00±8.66 b
Ae. polynesiensis
0.33±0.58 a
2.33±0.58 a
13.33±5.51 b
8.00±5.29 b
Cx. quinquefasciatus
2.33±1.53 a
1.00±0.00 a
8.00±4.36 b
3.67±1.53 ab
1
Means followed by the same letter are not significantly different (P<0.05); Tukey’s multiple comparison test applied
to log(x+1) transformed data.
afternoon and a lesser peak in the early morning, and
maximum biting activity was recorded between 1500 and
1800 h in both cases. Culex quinquefasciatus, however,
is generally accepted to be nocturnally active and
typically bites after 2200 h (Lee et al. 1989), although
early evening biting activity has been reported for the
Pacific from Fiji (Rakai et al. 1974).
Reports of mosquito collections using these
miniature light traps with various attractants have been
variable with respect to relative numbers and particular
species. Smith et al. (1979), in the initial comparison of
the newly developed EVS trap (Rohe and Fall 1979) with
the earlier CDC trap (Sudia and Chamberlain 1962), found
the former better for collecting Culex (Culex) tarsalis
Coquillett but not Culex (Culex) erythrothorax Dyar.
Later, Ritchie and Kline (1995) showed in southeast
Queensland, Australia that CDC-type traps (baited with
dry ice) consistently collected greater overall numbers
of mosquitoes than did EVS-type CO2 traps, but not for
all species. However, comparing collection data between
studies using different traps, and from different studies
using ostensibly the same traps, is problematic as even
relatively similar miniature light traps such as the CDC
and EVS traps differ in their component and structural
details. Moreover, the EVS-type used in the present study
was an advanced model, different than those used by
Ritchie and Kline (1995), and had more batteries (three
rather than two D-cell) driving an improved motor that
collected through an enlarged (11.5 cm rather than 8 cm
diameter) pipe section. The CDC-type used in the present
study was the “standard” model, with a 6-volt battery
and pipe diameter of 8 cm. The motors in the traps of the
present study were identical, but the propeller in the
CDC trap was four-bladed rather than two-bladed as in
the EVS trap, and the CDC trap lamp was a substantially
greater light source than that in the EVS trap.
The supplementation of carbon dioxide bait with
octenol has been reported to significantly increase
collections of some mosquitoes (including various aedine
species), but not others (particularly Culex species), in
various traps in various geographic regions (Takken and
Kline 1989, Kline et al. 1991, Kemme et al. 1993, Kline
Table 3. Numbers of Aedes aegypti, Aedes polynesiensis, and Culex quinquefasciatus collected in CDC- and EVStype light traps baited with carbon dioxide (dry ice) and also octenol (1-octen-3-ol), over 24 h for four d, in three
locations between Cooks Bay and Opunohu Bay, Moorea, French Polynesia, November 2003.
Trap type and bait
Species
Location
CDC
CO21
Ae. aegypti
EVS
CO2 + octenol
CO2
CO2 + octenol2
Pao Pao
14
9
18
11
Gump Stn
3
0
6
2
Vaihara
5
3
9
2
Ae. polynesiensis
Pao Pao
6
0
10
3
Gump Stn
26
4
19
18
Vaihara
31
7
18
7
Cx. quinquefasciatus
Pao Pao
15
2
19
3
Gump Stn
11
2
8
6
Vaihara
11
0
15
3
1
At Vaihara, Culex (Culex) roseni and Aedes (Aedimorphus) nocturnus were also captured - the latter being a new
species record for Moorea and French Polynesia. 2At Vaihara, Aedes (Aedimorphus) nocturnus was also captured.
December, 2004
Journal of Vector Ecology
313
Table 4. Mean number (± S.D.) of Aedes aegypti, Aedes polynesiensis and Culex quinquefasciatus collected in
CDC- and EVS-type light traps baited with carbon dioxide (dry ice) and also octenol (1-octen-3-ol) at Moorea,
French Polynesia, November 2003.
CDC
EVS
CO2
CO2+OCT
CO2
CO2+OCT
Ae. aegypti
7.33±5.85 a1
4.00±4.58 a
11.00±6.25 a
5.00±5.20 a
Ae. polynesiensis
21±13.23 a
3.67±3.51 a
15.67±4.93 a
9.33±7.76 a
Cx. quinquefasciatus
12.33±2.31 a
1.33±1.15 b
14.00±5.56 a
4.00±1.73 ab
1
Means followed by the same letter are not significantly different (P<0.05); Tukey’s multiple comparison test applied
to log(x+1) transformed data.
1994, Van Essen et al. 1994, Ritchie and Kline 1995). In
a comparison of octenol and carbon dioxide attracting
Ae. aegypti to Fay-Prince traps in Queensland, octenol
was reported to significantly decrease collections
(Canyon and Hii 1997). Similar results were reported
recently with another Stegomyia mosquito, where
significantly more Aedes (Stegomyia) albopictus (Skuse)
were collected in Maryland with Fay-Prince traps baited
with carbon dioxide or carbon dioxide plus octenol, when
compared with unbaited or octenol-only baited traps, and
octenol was found to be of no benefit (Shone et al. 2003).
However, there is no published information on the
reaction of Aedes (Stegomyia) species in the Pacific
region to CDC or EVS traps baited with carbon dioxide
and/or octenol. The data reported from the present study
indicate the species respond well to the use of carbon
dioxide in these traps, but there appears to be no value
in the use of octenol as an attractant, either alone or in
conjunction with carbon dioxide. Overall, however, the
numbers collected by either the CDC or EVS traps when
baited with carbon dioxide were not substantial when
compared with the standard technique of human bait
collections. This finding was similar to those reported
by Canyon and Hii (1997) who compared Fay-Prince traps
with human bait collections in Queensland, and Jones et
al. (2003) who found that human bait collections in
Thailand were much more effective than the omnidirectional Fay-Prince trap, the Centers for Disease
Control Wilton trap, and the Austech International “sticky
lure” in collecting Ae. aegypti adults for population
surveillance. Schoeler et al. (2004) reported from Peru
that the American Biophysics (ABC-PRO) light trap, the
omni-directional Fay-Prince trap, and the CDC Wilton
trap were not effective collectors of Ae. aegypti when
compared with human landing collections and
particularly with backpack-aspirator collections.
In conclusion, it would appear that human bait
landing/biting collections may be required to serve as
the most effective method for collecting adult Aedes
aegypti and other Stegomyia species in the Pacific region
for the foreseeable future.
Acknowledgments
This paper is contribution 114 of the University of
California Berkeley’s Richard B. Gump South Pacific
Research Station, Moorea French Polynesia. Financial
support was provided in part by a Gump Station Senior
Research Fellowship to the author who is grateful also
to Neil Davies (Executive Director of the Gump Station)
for providing facilities and to his staff, particularly Frank
Murphy and Valentine Brotherson, for assistance with
logistics and supplies. The author is also grateful to
Yves Sechan and Anne-Marie Legrand of the
entomology/filariasis team of the Institut Louis Malarde
(Papeete, Tahiti) for advice on local situations, Beth and
Victor Wong and Carol Raydon at Pao Pao/Cooks Bay,
and Tony You Sing at Vaihara/Opunohu Bay for allowing
collections at their homes, and Scott Ritchie and Cameron
Webb for statistical assistance and manuscript review.
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