Page 1
Size, shape and hue modulate attraction and landingresponses of the braconid parasitoid Fopius arisanus to fruitodour-baited visual targets
Jeanneth Perez • Julio C. Rojas • Pablo Montoya •
Pablo Liedo • Francisco J. Gonzalez • Alfredo Castillo
Received: 25 June 2011 / Accepted: 9 October 2011 / Published online: 16 November 2011
� International Organization for Biological Control (IOBC) 2011
Abstract Female parasitoids are guided by multi-
sensory information, including chemical and physical
cues during host location. In the present study, we
investigated the behavioural responses of naıve Fopius
arisanus (Sonan) females to visual targets baited with
guava odour. In non-choice wind tunnel tests, the
attraction and landing responses of parasitoids to
spheres painted with different colours, and targets of
different shapes and sizes were evaluated. Females
were more frequently attracted and landed more often
on dark yellow targets than on targets with other
colours. There was no correlation between the bright-
ness of each colour and the attraction or landing
responses. In contrast, both responses were correlated
with relative reflectance (hue) of the coloured targets.
A positive correlation was observed between attraction
and hue, and a negative correlation between landing
and hue. F. arisanus was attracted to and landed more
often on spheres than on other shape models. The
attraction response of this parasitoid was affected by
the size of the targets, with spheres of 10 and 12 cm
diameter being more attractive than spheres of 8, 6 and
4 cm diameter. The fact that F. arisanus females were
able to discriminate among visual targets that differ in
colour, shape and size stresses the importance of vision
during host location by this species.
Keywords Visual cues � Spectral reflectance �Attraction � Landing � Host location � Braconidae
Introduction
The host selection by parasitoids is a sequential
process that can be divided in host habitat location,
host location, and host acceptance (Vinson 1976;
Fatouros et al. 2008). At long-range, parasitoids forage
between plants used by their host within their habitat,
Handling Editor: Torsten Meiners
J. Perez (&) � J. C. Rojas � P. Liedo � A. Castillo
El Colegio de la Frontera Sur (ECOSUR), Carretera
Antiguo Aeropuerto km 2.5, 30700 Tapachula, Chiapas,
Mexico
e-mail: [email protected]
J. C. Rojas
e-mail: [email protected]
P. Liedo
e-mail: [email protected]
A. Castillo
e-mail: [email protected]
P. Montoya
Subdireccion de Desarrollo de Metodos, Programa
Moscas de la Fruta DGSV SAGARPA, Central Poniente
14, 30700 Tapachula, Chiapas, Mexico
e-mail: [email protected]
F. J. Gonzalez
Coordinacion para la Innovacion y la Aplicacion de la
Ciencia y la Tecnologıa, Universidad Autonoma de San
Luis Potosı, Sierra Leona 550, Lomas 2da. Seccion, 78210
San Luis Potosı, SLP, Mexico
e-mail: [email protected]
123
BioControl (2012) 57:405–414
DOI 10.1007/s10526-011-9416-0
Page 2
at medium-range they search for the hosts within host
plants, and finally at short-range females recognize
and accept their hosts (Volkl 2000). During the host
selection process, parasitoids are guided by multisen-
sory information, including chemical, vibratory and
visual cues, which are used in an interactive manner.
Chemical cues seem to play an important role in the
different steps within the host selection process,
whereas visual cues are thought to be important only
after females have been stimulated by host odours
(Vinson 1976).
The role of chemical cues during the host selection
process has been studied extensively in parasitoids,
but the use of vision during this process is less
understood for this group of insects. Visual informa-
tion may consist of variables such as colour, shape,
and size, among others. These parameters have
different behavioural relevance during host habitat
location at distance, and during host location after the
parasitoid has landed on a plant. At long-range, colour
seems to be the most important visual parameter,
while at short-range, shape and size are crucial for host
recognition and host acceptance (Wackers and Lewis
1999). Colour can be described in terms of three main
attributes: hue (dominant wavelength of reflected
light), brightness (intensity of perceived reflected
light) and saturation (spectral purity of reflected light)
(Prokopy and Owens 1983). True colour vision in
insects has only been confirmed for a limit number of
orders, among them Hymenoptera (Chittka and Men-
zel 1992; Kelber et al. 2003). Most of hymenopteran
species investigated so far have trichromatic vision
with UV (340 nm), blue (430 nm), and green
(535 nm) receptors (Peitsch et al. 1992). The capacity
for colour discrimination has been reported for several
species of parasitoids (Messing and Jang 1992;
Battaglia et al. 2000). For example, colour plays an
important role during host recognition and oviposition
by the parasitoid Aphidius ervi Haliday (Battaglia
et al. 2000). In addition to colour, other visual
parameters may be used by parasitoids during host
location, including the size and shape (Vinson 1976).
Host size and shape may be visual cues highly
distinguishable in almost any habitat because mor-
phological differences exist not only between plants
but also between structures of the same plant (Wackers
and Lewis 1999). The use of these parameters during
host location has been documented for different
parasitoid species. For instance, Diachasmimorpha
longicaudata (Ashmead) females showed no prefer-
ence for colour or shape, but displayed a clear
preference for large over smaller fruit-mimicking
models (Segura et al. 2007).
Fopius arisanus (Sonan) is an egg-pupal endopar-
asitoid that attacks several species of tephritid fruit
flies, particularly those in the genus Bactrocera.
Currently, F. arisanus is known to parasitize 21 fruit
fly species, and can develop in at least 18 species
(Quimio and Walter 2001; Rousse et al. 2005, 2006).
The successful introductions of this parasitoid to
Hawaii and French Polynesia for the control of
Ceratitis capitata (Wiedemann) and Bactrocera dor-
salis (Hendel), respectively, suggest that F. arisanus
has potential for augmentative biological control of
fruit flies, alone or in combination with other control
methods (Harris et al. 2000).
Host location behaviour of F. arisanus has been
described by Wang and Messing (2003). Chemical
cues play important roles during host location behav-
iour in this wasp (Liquido 1991; Altuzar et al. 2004;
Rousse et al. 2007a). However, there are contrasting
reports regarding the role of visual preferences during
host location (Vargas et al. 1991; Cornelius et al. 1999;
Rousse et al. 2007b). With the aim of clarifying the
role of visual preferences in host location by F.
arisanus, this study was undertaken to investigate the
behavioural responses of female wasps to guava
odour-baited targets of different colours, sizes and
shapes in a laboratory wind tunnel.
Materials and methods
Insects
Parasitoids used in this study were obtained from
colonies established at MOSCAFRUT mass-rearing
facility (SAGARPA-IICA) located in Metapa de
Domınguez, Chiapas, Mexico. This strain of parasit-
oids was imported from Hawaii in 1998, where it was
reared on C. capitata eggs (Bautista et al. 1999).
Parasitoids used for experiments had been reared from
Anastrepha ludens (Loew) eggs placed in pieces of
papaya at 22 ± 2�C, 60–80% RH, and 12:12 L:D h
photoperiod as previously described (Montoya et al.
2009). The eggs of A. ludens were obtained from adults
reared on artificial diet according to established
protocols (Domınguez et al. 2010). After emergence,
406 J. Perez et al.
123
Page 3
parasitoids of both sexes were placed in wooden frame
cages 30 9 30 9 30 cm to allow mating. Adults were
given free access to honey and water dispensed on
cotton wool. One hour before the experiments, females
were placed individually in plastic containers (5 cm
high by 4 cm diameter) and allowed to acclimatize to
the wind tunnel room conditions. In all experiments,
7–9 days old females were used. Parasitoids had no
prior contact with their hosts or host fruits.
Bioassays
The observations were performed in a wind tunnel
(120 cm long 9 30 cm high 9 30 cm wide). The
wind tunnel was constructed using 1 cm thick clear
plexiglass. A fan was used to pull air, filtered by
activated charcoal, through the tunnel at a speed of
0.4 m s-1. Illumination was provided by four fluores-
cent tubes mounted 60 cm above the wind tunnel
giving a light intensity of 2380 lx. Green dots of
10 cm diameter made from cardboard were placed on
the tunnel floor to provide optomotor feedback for the
parasitoids. Fruit odour consisted of five green guavas
(325.7 ± 13.2 g) (mean ± SD) without insect infes-
tation, placed inside a polyethylene bag that was
situated outside the wind tunnel. Filtered air
(1 l min-1) was passed through the bag and the odour
was conducted into the wind tunnel by means of tygon
tubing (0.4 cm internal diameter). The end of the tube
was connected to a hole (0.7 cm) in the centre of the
visual target from which odour emanated at a flow rate
of 0.6 l min-1. The target stimulus was hung in the
centre of the wind tunnel, 16 cm from the upwind end.
The distance between platform release and the target
stimulus was about 1 m.
Each observation started by placing the plastic
container with one female on a 12 cm high platform at
the downwind end of the tunnel. The parasitoid was
released and observed for 5 min. Female behaviour
was recorded, specifically the zigzagging upwind
flight towards the target stimulus and landing on the
target after such upwind flight. If the parasitoid did not
take off or did not show zigzagging upwind flight after
5 min, the test was stopped and the insect was
considered to be a non-responding individual. Here-
after, the term attraction is used to indicate such
zigzagging upwind flights. Parasitoids were used
once. In all experiments, the responses of parasitoids
to the visual targets were evaluated in non-choice
tests. All bioassays were conducted between 08:00 h
and 14:00 h at 24 ± 2�C, 65 ± 5% RH.
Response to models of different colours
The responses of F. arisanus females to styrofoam
spheres of different colours baited with guava odour
were evaluated. Preliminary observations showed that
females were not attracted to spheres without the
odour. Spheres of 8 cm diameter were painted with the
following colours of acrylic paint (Comex Group,
Mexico): white, black, brown, red, blue, light green,
dark green, light yellow and dark yellow. The spectral
reflectance curves of the different colours were mea-
sured using a spectrometer (USB4000-VIS-NIR,
Ocean Optics, Dunedin, FL, USA) with an optical
resolution of *1.5 nm (full width at half maximum), a
tungsten-halogen light source (LS-1, Ocean Optics,
Dunedin, FL) and a reflection probe (R200-7-VIS-NIR,
Ocean Optics, Dunedin, FL, USA). The raw reflectance
spectra were corrected for detector dark current and
normalized to the spectrum obtained from the light
source reflected on a white reference standard (Fig. 1).
Five replicates per colour were made daily in a random
order and the experiment was repeated on different
days until completing the 45 replications per colour.
Response to models of different shape
We evaluated the responses of F. arisanus females to
models of different shapes with similar surface area
Wavelength (nm)400 500 600 700
Spec
tral
ref
lect
ance
(%
)
0
3
6
20
40
60
80W
Dy
R
LyB
Dg
BrLgBl
Fig. 1 Spectral reflectance curves of the nine colours used in
the experiment Dy dark yellow, Ly light yellow, B blue, W white,
Br brown, Bl black, R red, Lg light green and Dg dark green
Responses of the braconid 407
123
Page 4
(313 cm2). All models were made using styrofoam
and were painted dark yellow. This colour was chosen
because it was the most attractive to F. arisanus
females in the previous experiment. The models
evaluated had either three-dimensional (sphere, cube,
pyramid, and ellipse) or flat shapes (circle, square,
rectangle, and triangle). Five replicates per shape were
performed daily in a random order and the experiment
was repeated on different days until completing 45
replicates per shape.
Response to models of different size
The responses of F. arisanus females to dark yellow
spheres of 14, 12, 10, 8, 6, and 4 cm diameter were
tested in this experiment. The size of spheres were
chosen on the basis of previous studies with this
species and considering the range of size of fruits used
by F. arisanus hosts. Five replicates per size were
made daily in a random order and the experiment was
repeated on different days until completing 45 repli-
cates per size.
Statistical analysis
We used a log likelihood ratio contingency-table
analysis to test whether responses (attraction or
landing) differed among treatments. Fisher’s exact
test was used when counts were low. When significant
effects were found, odds ratios were used to make
comparisons between treatments.
In the first experiment, numbers of attraction and
landing responses were correlated with the brightness
and hues of visual targets, using methods described
elsewhere (Katsoyannos and Kouloussis 2001).
According to this procedure whether the insect
response to coloured target depends only on bright-
ness, a strong correlation between the frequency of
responses and total brightness was expected. If the
insect response depends on the specific wavelengths
(hues), then the response should correlate with relative
reflectance values of the colours. Relative reflectance
was considered to be the amount of light reflected by a
colour within a given wavelength interval relative to
the total amount of light reflected by this colour within
the range of the spectrum considered.
The total reflectance values of the colours used in this
study were obtained as percentages of the total amount
of light reflected over a white reference sphere between
350 and 750 nm. Reflectance values were: white =
100, dark yellow = 51, light yellow = 14.1, dark
green = 7.9, light green = 9.7, blue = 9, brown =
8.9, black = 7.1, and red = 24. Relative reflectance
were calculated for every colour as the amount of light
reflected by each successive 10 nm wide wavelength
interval, in relation to the total amount of light reflected
by that colour between 350 and 750 nm.
Results
Response to models of different colours
Colour influenced the attraction of F. arisanus females
towards the spheres baited with fruit odour (G = 21.1,
df = 8, P = 0.007). Females were more attracted to
dark yellow models than to light yellow, blue, black,
red, light green, and dark green models (Fig. 2a).
There were no differences in the numbers of females
that were attracted to dark yellow, white, and brown
targets. The landing behaviour of parasitoids on the
targets was also influenced by colour (G = 67.44,
df = 8, P \ 0.001). Females landed more often on the
dark yellow, light yellow, brown, black, red, and light
green models than on blue, white and dark green
targets (Fig. 2b).
There was no correlation between the total amount
of reflected light of each colour and attraction
(r = 0.44) or landing (r = -0.399) responses (P [0.05). In contrast, both responses were correlated with
relative reflectance (hues) of the coloured targets. A
positive correlation was detected between attraction
and hue, and a negative correlation between landing
and hue (Fig. 3). Females were more attracted to hues
reflecting maximally between 550 and 620 nm (dark
yellow), with a maximum peak at 600 nm (r = 0.795).
Parasitoids landed less often on hues reflecting
maximally between 420 and 510 nm (blue, light
green), with a minimal peak at 490 nm (r = -0.943).
Response to models of different shape
The shape of visual targets affected attraction
(G = 24.2, df = 7, P = 0.001) and landing (G =
19.0, df = 7, P = 0.008) responses of F. arisanus.
Parasitoids were more attracted to spheres than to
square, rectangle, or pyramid models. Attraction of
F. arisanus females to spheres was not significantly
408 J. Perez et al.
123
Page 5
different from that of circle, ellipse, cube and triangle
models (Fig. 4a). Parasitoids landed more often on
sphere, circle, and cube models than on the triangle
model. The landing response on ellipse, square,
rectangle and pyramid were intermediate and not
significantly different from that on sphere, circle and
cube or triangle (Fig. 4b).
Response to models of different size
The size of the visual targets affected attraction of
F. arisanus females (G = 33.7, df = 5, P \ 0.001).
In contrast, the landing response was not affected by
the size of the models tested (Fisher test, P = 0.76).
Parasitoids were more attracted to spheres of 10 and
12 cm diameter than to spheres of 4, 6 and 8 cm
diameter. The attraction of F. arisanus to spheres of
14 cm diameter was intermediate and not significantly
different from that to spheres of 10 and 12 cm
diameter (Fig. 5).
Discussion
We showed that some visual cues in combination with
fruit odour might play a role in the host location
0
20
40
60
80
100
DarkYellow
LightYellow Green
DarkGreen
Att
ract
ion
resp
onse
(%
)
Colour
bc
a
bc bcbc
bc
c
ab ab
a
0
20
40
60
80
100
DarkYellow
LightYellow
Blue White Brown Black Red Light
Blue White Brown Black Red LightGreen
DarkGreen
Lan
ding
res
pons
e (%
)
Colour
a
a a
aa
a
b
b
b
b
Fig. 2 Attraction (i.e.
zigzag upwind flight
towards the visual target
stimulus) (a) and landing
(b) (±S.E.) of F. arisanusfemales on visual targets of
different colour. Bars
capped with the same letter
are not significant different
(P [ 0.05)
Responses of the braconid 409
123
Page 6
behaviour of naıve F. arisanus females. We found that
females of this parasitoid were attracted and landed
more often on yellow spheres compared with spheres
of other colours. Previously, Vargas et al. (1991)
reported that yellow and orange traps captured more
F. arisanus than those caught by blue and black traps.
Rousse et al. (2007b) showed that naıve F. arisanus
females landed less often on yellow and white spheres
than on the black and dark green spheres. These
authors concluded that F. arisanus females respond
mainly to achromatic cues. However, the response that
we obtained towards the dark yellow and brown
spheres showed that they were equally attractive to
F. arisanus, although they differ widely in the amount
of reflected light, suggesting that for this species the
hue is more important than brightness. An explanation
for differences between both studies may be due to the
experimental protocols. In our case, experiments were
performed using fruit odour in combination with
visual targets, and this may have affected the parasit-
oid responses. Thus, we consider that the response of
F. arisanus females to odour-baited visual targets may
be related to host foraging, while the responses of the
parasitoids reported by Rousse et al. (2007b) may only
imply the ability of females to detect and respond to
visual targets, without direct link to the host location
process. On other hand, our bioassays were performed
in a wind tunnel with controlled illumination, such that
the amount of light received by the coloured visual
targets of the different treatments was constant
throughout the whole experiment. Rousse et al.
(2007b) performed experiments in field cages where
the amount of light received by each visual target
likely varied depending on their position relative to the
sun and the target, and variation in solar and clima-
tological condition, thus influencing the parasitoid’s
responses. Indeed, the received colour of an object
depends upon the interaction between the ambient
light and the reflectance colour of the object. Thus, an
object may have a different appearance in each
environment (Endler 1993). Another reason is that in
our study the visual targets were not placed against a
particular background (black, white, gray). Consider-
ing that insects likely perceive foliage as gray, thus
against this achromatic background, any different
wavelength other than that of foliage will produce at
least some colour contrast (Chittka et al. 1994).
The preference for the colour yellow we found in
F. arisanus has been reported for other hymenopterans,
including egg parasitoids (Lobdell et al. 2005). In the
case of fruit fly parasitoids, Diachasmimorpha juglan-
dis (Muesebeck) chose yellow walnut fruits over black
fruits despite that the latter are more likely to contain
host larvae (Henneman 1998). In field cage tests,
D. longicaudata females discriminated between vari-
ous spheres painted with different colours with yellow
as the most attractive colour (Messing and Jang 1992).
In the same way, several studies have shown that
yellow colour is highly attractive to different species of
fruit flies (Katsoyannos 1987; Vargas et al. 1991; Drew
et al. 2003; Pinero et al. 2006; Lopez-Guillen et al.
2009). For example, C. capitata females were mainly
attracted to yellow, followed by red, orange, black, and
green spheres, whereas white and blue spheres were the
least attractive (Katsoyannos 1987).
The shape and size of targets are other visual
parameters that can be used during host location by
parasitoids. The hue and brightness of visual targets
can be perceived before insect landing, whereas the
shape and size gain in importance at lesser distances.
Vinson (1976) pointed out that size and shape are
usually secondary stimuli during foraging, and
become important during host acceptance. However,
in some cases these stimuli seem to be crucial during
host location. For instance, Henneman et al. (2002)
suggested that at distance, D. juglandis females tend to
ignore the odour and use visual cues, because the
odour plume is turbulent and difficult to locate the host
fruit. We have found that F. arisanus females showed
better responses to spherical than to rectangular or
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
1.0
350 390 430 480 520 560 600 650 690 740
Cor
rela
tion
coe
ffic
ient
(r)
Wavelength (nm)
Attraction Landing
P < 0.05
P < 0.05
Fig. 3 Correlation coefficient between the responses (attrac-
tion and landing) of F. arisanus females to targets of nine
different colours, and the relative reflectance values of each
colour at successive 10 nm wavelength, from 350 to 750 nm.
Statistical differences are indicated above or below of the dash-
lines
410 J. Perez et al.
123
Page 7
square models. It is important to mention that we
performed this bioassay using yellow models only.
Therefore, further experiments are needed to investi-
gate whether shape may interact with colour as it is
documented for other insects (Bernays and Chapman
1994). Also, we found that parasitoids were more
attracted to spheres between 10 and 14 cm diameter.
Diachasmimorpha longicaudata females also showed
preferences for larger spheres (Segura et al. 2007).
Similarly, the shape and size of visual targets influence
host selection in different species of fruit flies. Adults
of C. capitata and Anastrepha fraterculus (Wiede-
mann) are more attracted to spherical forms than to
rectangular models (Cytrynowicz et al. 1982). Lopez-
Guillen et al. (2009) reported that spherical models
captured more Anastrepha obliqua (Macquart) adults
than those caught by rectangular, triangular, or
circular models. Also, they observed that spheres of
8, 10, 12 cm diameter captured more A. obliqua adults
than smaller spheres. It has been suggested that fruit
flies are more attracted to spherical models because
this shape mimics the fruits used by insects for
oviposition (Katsoyannos 1989). A similar idea may
be applied to explain the results obtained with
F. arisanus females, where colour, size and shape
could have a significant influence in host location.
0
20
40
60
80
100
Att
ract
ion
resp
onse
(%
)
Shape
a
abab
abc
d
cd
abc
bcd
a
0
20
40
60
80
100
Sphere Circle Ellipse Cube Square Rectangle Triangle Pyramid
Sphere Circle Ellipse Cube Square Rectangle Triangle Pyramid
Lan
ding
res
pons
e (%
)
Shape
b
ab
a
ab
ab
ab
bb
b
Fig. 4 Attraction (i.e.
zigzag upwind flight
towards the visual target
stimulus) (a) and landing
(b) (±S.E.) of F. arisanus on
visual targets of different
shape. Bars capped with the
same letter are not
significant different
(P [ 0.05)
Responses of the braconid 411
123
Page 8
Insects usually use multisensory information during
foraging (Harris and Foster 1995). In this way, Hebets
and Papaj (2005) proposed several hypotheses con-
cerning the role of multimodal signals, including the
possibility that a combination of cues facilitates the
detection of a target at different distances, where the
detection of one sensory modality alerts the insect to
the presence of the other. In this sense, we used fruit
odour in all our experiments because parasitoids were
not attracted to visual targets when they were offered
alone. Our results emphasize the importance of the
interaction of different cues during host selection by
this parasitoid. Thus, although we used the same
olfactory cue, females were able to discriminate
among the different models offered indicating the
importance of visual cues at long-range. The impor-
tance of visual cues at short-range was also observed
in this study. For example, although the white colour
was as attractive as the dark yellow, only 36.6% of
parasitoids that were attracted to white spheres landed
on them. The remaining females that did not land on
white spheres (23%) hovered at 1–2 cm from the
target for about 3 min, and then returned to the
downwind end of the tunnel, sometimes landing on the
release platform. This result is in agreement with the
idea that vision has a critical role in the final stages of
host location, partly because the lack of odour
gradients makes it most improbable that odour would
guide an insect directly to a plant, but also because a
flying insect needs to use vision during landing
(Bernays and Chapman 1994). The multimodal sen-
sory integration has been reported for other parasitoid
species as well. Jang et al. (2000) suggest that during
foraging, visual and olfactory cues, both alone and in
combination, affect the behaviour of D. longicaudata
females. Likewise, Jonsson et al. (2005) reported an
increase preference for flower odour when colours
were added in the bioassays performed with the
ichneumonid females of Phradis interstitialis Thom-
son and Tersilochus heterocerus Thomson, parasitoids
of Meligethes spp. (Coleoptera: Nitidulidae). Finally,
the tachinid parasitoid Exorista japonica Townsend
uses both visual and olfactory cues to locate the host
habitat of the noctuid pest Mythimna separata
(Walker) (Ichiki et al. 2011).
Our results suggest that F. arisanus attraction to
coloured targets is due to the hue rather than the
intensity of reflected light. We have showed that
F. arisanus can discriminate between colours, but
without odours these visual cues mean nothing for
naıve F. arisanus females, at least within the limits of
our bioassays. From this, we infer that females are
initially stimulated by fruit odours and are then guided
to targets by a combination of visual and chemical
cues. The fact that parasitoids were able to discrim-
inate among visual targets that differed in colour,
shape and size, stresses the importance of vision
during host location by this species.
Acknowledgments We are grateful for the technical assistance
of Florida Lopez (PROGRAMA MOSCAFRUT) and Armando
Virgen (ECOSUR). Rosalba Morales provided logistic support.
Many thanks go to Javier Valle Mora for statistical advice and
Francisco Infante and Trevor Williams for a critical review of a
previous version of the manuscript. J. Perez gives special thanks to
El Consejo Nacional de Ciencia y Tecnologıa (CONACYT) of
Mexico for the doctoral scholarship.
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0
20
40
60
80
100
14 12 10 8 6 4
Att
ract
ion
resp
onse
(%
)
Diameter (cm)
bc
aab
a
cc
Fig. 5 Attraction (i.e. zigzag upwind flight towards the visual
target stimulus) (±S.E.) of F. arisanus on visual targets of
different size. Bars capped with the same letter are not
significant different (P [ 0.05)
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Author Biographies
Jeanneth Perez Received her MSc degree from El Colegio de
la Frontera Sur, Mexico. At present she is a Ph. D candidate at
the same institution. Her research interests are chemical
ecology, insect behaviour and biological control of tropical
insects.
Julio C. Rojas Received his PhD from Oxford University in
1998. He is a research entomologist at the Department of
Tropical Entomology, El Colegio de la Frontera Sur, Mexico.
His interests include the behaviour and the chemical ecology of
herbivorous and carnivorous insects.
Pablo Montoya Doctor in Sciences from the Universidad
Nacional Autonoma de Mexico in 1999. He is a Sub-Director
and researcher in the Moscafrut Program SAGARPA-IICA of
Mexico. He has experience in the management of fruit flies,
particularly in biological control, detection of populations, and
the application of the Sterile Insect Technique.
Pablo Liedo Ph. D. in Entomology, University of California -
Davis (1989). He is a research entomologist at the Department
of Tropical Entomology, El Colegio de la Frontera Sur,
Mexico. His area of interest is insect ecology and pest control,
with particular emphasis on biodemography, population ecol-
ogy, biological control and the Sterile Insect Technique.
Research work has been focused on fruit flies.
Francisco J. Gonzalez Ph.D. in electrical engineering from the
School of Optics and Photonics, University of Central Florida,
Orlando, in 2003. He is currently a Professor at the Universidad
Autonoma de San Luis Potosı, Mexico. His research interests
are the application of physics, optics and engineering in non-
invasive medical diagnosis.
Alfredo Castillo Doctor in Science from El Colegio de
Posgraduados in 2005. He is an entomologist at the Department
of Tropical Entomology, El Colegio de la Frontera Sur,
Mexico. He conducts research on ecology and biology of
tropical insects.
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