J Insect Behav

DOI 10.1007/s10905-012-9325-9

Male Wing Vibration in the Mating Behavior

of the Olive Fruit Fly Bactrocera oleae (Rossi)
(Diptera: Tephritidae)
Giovanni Benelli & Angelo Canale &
Gabriella Bonsignori & Giacomo Ragni &
Cesare Stefanini & Alfio Raspi

Revised: 1 March 2012 / Accepted: 6 March 2012

# Springer Science+Business Media, LLC 2012

Abstract The olive fruit fly, Bactrocera oleae (Diptera: Tephritidae), is a worldwide
pest of olive fruits. To date, the mating behavior of this tephritid has not been
fully clarified. Little has been reported regarding the characteristics and behavioral role of B. oleae male wing vibration. This study provides a detailed description
of male wing vibration, recorded by a high-speed video camera and compared with a
frame-by-frame analysis, prior to successful and unsuccessful matings. Experimental
evidence is given of the importance of male wing vibration during courtship for
successful mating. Firstly, when the males did not perform wing vibration, but simply
jumped onto the female and immediately attempted to copulate, no successful
matings were registered. Secondly, when the males wings were removed, the probability of successful mating was significantly reduced. Thirdly, the frequency and
pulse duration of the male wing vibration during courtship were higher in successful
matings than in unsuccessful ones. Lastly, a wing sexual dimorphism was found:
length and total wing area were significantly higher among males. Overall, our
research increases the knowledge of the reproductive behavior of B. oleae, by
suggesting the importance of wing vibration among the range of sensory modalities
used by this tephritid in sexual communication.
Keywords Courtship . Dacinae . fruit flies . intra-sexual communication . mating cues

G. Benelli : A. Canale (*) : A. Raspi

Department of Tree Science, Entomology and Plant Pathology, University of Pisa,
via San Michele degli Scalzi 2, 56124 Pisa, Italy
e-mail: acanale@agr.unipi.it
G. Bonsignori : G. Ragni : C. Stefanini
BioRobotics Institute, SantAnna School of Advanced Studies, viale Rinaldo Piaggio 34,
56025 Pontedera, Italy

J Insect Behav

Introduction
Among Diptera, acalyptrate flies use many different communication cues in during
courtship and mating (Burk 1981). Behavioral studies - conducted mainly under
laboratory conditions - have shown that many species of Tephritidae use visual,
auditory and chemical stimuli for their mating communication (Mazomenos 1989).
In several fruit fly species, courting males vibrate their wings and produce speciesspecific signals (Keiser et al. 1973; Webb et al. 1983; Sivinski et al. 1984). Wing
vibration has been studied intensively in the tephritids Ceratitis capitata (Briceo and
Eberhard 2000a; Briceo and Eberhard 2002; Briceo et al. 2002, 2007), Anastrepha
suspensa (Webb et al. 1984), Bactrocera cucurbitae (Miyatake and Kanmiya 2004),
Bactrocera dorsalis (Poramarcom 1988; Poramarcom and Boake 1991), and in
Drosophila (Colegrave et al. 2000 and references therein). In A. suspensa, male wing
vibrations increase female activity (Burk 1981) and play an important role both in
male-male competition during courtship (Sivinski et al. 1984) and in attracting
females (Webb et al. 1976). In the genus Drosophila, male wing vibrations affect
female receptivity (Kyriacou and Hall 1982; Aspi and Hoikkala 1995), inhibit female
locomotion, or increase sexual behavior in nearby males (von Schilcher 1976). Of the
various species of tephritids and drosophilids, the parameters of wing signaling may
vary according to geographical origin (Calcagno et al. 1999; Briceo et al. 2007),
strain (Colegrave et al. 2000; Briceo and Eberhard 2002), mass-rearing (Sivinski et
al. 1989; Briceo and Eberhard 2002), age (Miyatake and Kanmiya 2004), and diet
(Diamantidis et al. 2008).
The olive fruit fly, Bactrocera oleae (Rossi), is a tephritid pest of olives in many
countries of the Mediterranean basin (Daane and Johnson 2010). Recently, this
species was discovered in southern California and then spread widely throughout
the state, posing a serious threat to the olive industry (Rice et al. 2003). For
table olives the oviposition punctures lead to a serious reduction in crop value. In
olives cultivated for oil production the damage consists mainly of premature fruit
drop and a lower quality of the oil (Tzanakakis 2006). Female olive fruit flies are
oligogamous and mate 13 times during their lifetime (Tzanakakis et al. 1968;
Cavalloro and Delrio 1970; Zouros and Krimbas 1970). Male olive fruit flies are
polygamous and can mate daily if receptive females are available (Zervas 1982). In
laboratory and field conditions, matings occur in the late afternoon or at dusk,
indicating that there is a daily rhythm in sexual activities (Mazomenos 1989).
Olfactory cues from both sexes are known to be crucial in B. oleae courtship and
mating. In a laboratory study, Haniotakis (1974) proved that sexually mature virgin
females attracted males during the mating period, suggesting that virgin females
release an airborne sex pheromone causing a short and long range orientation of
males to females. These results were verified in field tests, with traps baited with
virgin females (Haniotakis 1977). Further research reported the chemical structure of
the major component of the sex pheromone, 1,7-dioxaspiro-[5.5]-undecane, produced
by glands associated with the rectal ampulla in B. oleae females (Baker et al. 1980).
On the other hand, De Marzo et al. (1978) showed that female olive fruit flies were
attracted to males in olfactometer tests. These authors suspected that the male olive
fruit fly produces a pheromone which facilitates mating and that the rectum is the site of
production. This latter hypothesis was recently reinforced by a study of Mavraganis et

J Insect Behav

al. (2010), which reported that extracts from B. oleae male bodies attract virgin
females.
Little is known regarding cues produced during courtship and mating behavior of
B. oleae males. During courtship, B. oleae males vibrate their wings and stridulate by
rubbing the wing microtrichia on the A1 +CuA2 vein against the abdominal pecten
located on the edge of the third abdominal tergite (Feron and Andriew 1962; von
Rolli 1976; Loher and Zervas 1979). Feron and Andriew (1962) interpreted wing
vibration as a mating stimulus that may serve to test the females willingness to
copulate. It was unclear if this signal could be generated by the air-borne mechanical
particle displacement due to wing vibration, by stridulation, or by both mechanisms.
Among other Bactrocera species, Keiser et al. (1973) speculated that wing vibration
could be displayed by males as a territorial behavior during mating. To date, the
characteristics and behavioral role of B. oleae male wing vibration have not been
studied in detail. We hypothesize that B. oleae male wing vibration could play a key
role in the courtship phase for successful mating. The aim of our research is thus to (i)
describe the olive fruit fly courtship and mating behavior, with special regard to
the effect of the mating status of both sexes on male courtship decisions, (ii)
characterize the male wing vibration and (iii) provide evidence of its critical role in
courtship and mating.

Materials and Methods

Insect Rearing
Insects used in this study were obtained from field-derived pupae (collected in a
Tuscan olive-mill) during December 2010. Pupae were maintained in Pisa laboratory
under controlled conditions (22C1, 50-60 % R.H. and natural photoperiod) to wait
for adult emergence. To obtain coeval virgin specimens, emergent flies were sexed
during the first 12 h of life and placed in different Plexigas cylindrical cages (diameter
40 cm, length 50 cm, 50 flies per cage). Adults were fed on a dry diet (yeast and sugar
at ratio 10:1) and water (Gen and Nation 2008).
General Observations
Each trial was conducted in a 12 m2 room, illuminated with daylight fluorescent
tubes. These tubes were placed in such a way as to guarantee that the intensity of light
was as even as possible. The temperature was set at 22C1 and the relative humidity
at 45 %5. Bioassays were all performed during January 120, 2011 between 15.30
and 17.30 h, with 912 day-old flies, since they were sexually mature (Mazomenos
1989). For each replicate, flies were replaced by new ones of the same age. The
assays were carried out over several days to account for any daily variability.
Courtship and Mating Behavior
To observe the courtship and mating behavior of B. oleae, a virgin focal male was
gently transferred in a cylindrical glass testing arena (diameter 30 cm, height 35 cm)

J Insect Behav

containing 15 virgin males and 15 virgin females, and its behavior was noted by an
observer for 30 min or until the end of the mating. Before the trial, the focal male was
cooled for 3 min at 10 C, then marked with a small dot of non-toxic color paint
(Polycolor 256, Maimeri, Italy) on the thorax (Dukas and Mooers 2003). Preliminary
assays revealed that this treatment did not influence the fly behavior. Flies courtship
and mating behavior was video recorded with an HotShot 512 SC high-speed video
camera (NAC Image Technology Inc, Simi Valley, CA, USA). Fifty replicates were
performed. To observe if the female mating status can affect the males behavior, a
virgin or mated (mating occurred about 48 h before) male was gently transferred in the
testing arena containing 30 virgin or mated females. The male behavior was
observed for 30 min, if mating did not occur before. The number of males that
performed wing vibrations and copulation attempts was recorded. Fifty replicates
were performed for each trial. Data were processed by JMP 7 (SAS), using a
weighted generalized linear model (GLM) with one fixed factor: y X Q " in
which y is the vector of the observations (the number of males that performed wing
vibration or copulation attempts), is the vector of fixed effects (female mating status:
virgin or mated), and is the vector of the random residual effects (Sprinthall 2003).
Wing Vibration Video Characterization
B. oleae courtship and mating behavior was recorded with the HotShot 512 SC
high-speed video camera described above. Sequential images from each detail were
captured at a rate of 8000 fps with an exposure time of 0.125 ms. The HotShot 512
SC video camera stores images with a resolution of 512x256 pixels directly to its
internal memory. Then, these images were downloaded into a dedicated computer for
data analysis. The area in which insects were expected to perform mating behavior was
lit with four LED illuminators (RODER SRL, Oglianico, TO, Italy) that emit light
(420 lm each) at l0628 nm. The red light was chosen, both because it matches the
maximum absorption frequency of the camera and because it does not damage the visual
apparatus of the insects, not possessing receptors for that wavelength (Briscoe and
Chittka 2001). A virgin male was gently transferred in the testing arena containing
15 virgin females.
Wing vibrations performed prior to successful and unsuccessful matings were
compared with a frame by frame analysis of videotapes. To avoid pseudoreplicates,
only a single wing vibration sequence was analysed for each male. For each wing
vibration, the following parameters were measured (Briceo et al. 1996): the angle ()
of male orientation toward the female and of alignment between male and female
during wing vibrations and prior to copulation attempt, the distance (mm) between
the two flies (based on prothorax to prothorax distance) during wing vibrations and
prior to the male contact the female, the total time (s) that the female remained
immobile before the males jump, the mean amplitude () of the wing beat, the mean
pulse frequency (Hz) (the inverse of the average duration of the wingbeat during a
pulse, measured with the frame by frame analysis (8000 fps) of video recordings), the
pulse duration (s) (the time from the beginning of one pulse to the end of the same
pulse), and interpulse interval (s) (the time from the beginning of a pulse to the start of
the next pulse). Following Joyce et al. (2010), for each male three interpulse intervals
of the first series of 15 or more consecutive interpulse intervals were used to

J Insect Behav

characterize male wing vibration during courtship and copulation attempts: the
second interpulse from the beginning of the series, the middle interpulse of the series,
and the second to the last interpulse at the end of the series. When there was an even
number of pulses in the series, the middle pulse was calculated using the formula
[(total pulses/2)+1]. For each male, data from three interpulse intervals were averaged across them. All averages were followed by standard deviation. Thirty males
were filmed both for successful and unsuccessful matings. Selected high-speed
videos were edited with NAC HSSC Link software (Copyright 20042007 NAC
Image Technology Inc, Simi Valley, CA, USA) and analyzed with ProAnalyst suite
(Copyright 20032008 Xcitex Inc, Cambridge, MA, USA). The variance between
values was analyzed with Fishers F-test. Studentst-test was used to evaluate the
statistical differences in mean values between different trials.
To verify if wing sexual dimorphism exists, images of 50 wings of both sexes were
mounted on microscope slides, captured with a Nikon D 300S equipped with a
Zeiss Tessovar photomicroscope, and imported into a computer with an Adobe
Photoshop CS2 software. Landmarks were used to estimate length and width and to
divide the wing into anterior and posterior portion (Fig. 5). Length, width, and areas
were measured using a made tool (Wing Analyzer), implemented in Matlab code
(Matlab R2011A). Data were analyzed with R version 2.11.2 (Copyright 2010 The R Foundation for Statistical Computing, Vienna, Austria). The variance between
values was analyzed with Fishers F-test. Studentst-test was used to evaluate the
statistical differences in mean values between male and female wings.
Role of Male Wing Vibration in Mating
A virgin - wingless or not - male was gently transferred in the testing arena containing
30 virgin or mated females and observed for 30 min. The occurrence of a successful
mating was noted. To obtain wingless specimens, males were immobilized at 10C
for 3 min, and each wing was cut off just above the base. Wingless males were tested
after a waiting period of 15 min from the wing removal. To establish whether the cold
treatment alone would affect the behavior of males, a set of controls was run first: 60
males were cold immobilized as above, but their wings were not cut off before
exposure. Sixty replicates were done for each trial. Number of matings were processed by JMP 7, using the weighted GLM described above, in which y is the vector
of the observations (the male mating success), is the vector of fixed effects (the
male status: wingless or not), and is the vector of the random residual effects.

Results
Courtship and Mating Behavior
The sequence of events leading to copulation in B. oleae is reported into Fig. 1. It can
be divided in three main phases: (1) an initial phase that culminated with the males
arrestment, in which visual and olfactory cues could play an important role, (2) a
close-range phase, with male wing vibrations, and (3) a final contact phase, with
copulation attempts and leg scrubs (i.e., the male rubbed periodically its legs on the

J Insect Behav

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Fig. 1

A flow chart of courtship and mating behavior of B. oleae (n050 couples). Percentages indicate the
proportion of individuals displaying each behavior. Latency phase: the time elapsed before walking activity
commenced. Arrestment: the time in which the male remained immobile before starting wing vibrations.
Male body rocking and intromission: the time elapsed between the males landing on the female and the
occurred intromission, with genitalia contact. The large dashed line evidenced how the courtship sequence can
be affected by other males. The dotted line shows the percentage of males that did not perform wing vibrations
and tried to copulate directly with the female. Mean values (s) are followed by the relative standard deviation

females thorax and abdomen), in which tactile cues probably dominated. When released,
most males remained still for some seconds (latency phase) before starting to walk, with
their wings hold over the abdomen. When they met a conspecific, they frequently walked
with their wings open (about 90 respect to the longitudinal axis of fly body) and tried to
push it away. When a female was detected, receptive males stop walking and remained
still. Then, the majority of males started wing vibrations near the female. During courtship,
B. oleae males maintained their wings close to the abdomen, vibrating them fast in a
dorso-ventral fashion and rolling them on their longitudinal axes (Fig. 2). When
performing wing vibration, the male wing posterior portion repeatedly contacted on
the abdomen, rubbing microtrichia placed around the A1 +CuA2 vein on the abdominal pecten. Some males did not court, but simply jumped on the female and
immediately attempted to copulate. No successful matings were observed in these
males. Moreover, if the mating sequence was interrupted by another male, the first
male was frequently able to resume courtship of the same female. Time spent in
precopulatory activities was 589.9386.49 s. In attempting copulation, the male
jumped onto the female from the back or side, trying to align himself along her body
axis. When receptive, the female remained still, allowing males intromission. During
Fig. 2 Lateral (a, b, c)
and frontal (a1, b1, c1) view
of B. oleae male during wing
vibration. When vibration starts,
the wings are close to the
abdomen and approximately
parallel to the ground (a, a1).
Then, the wings are rapidly
raised and rolled on their
longitudinal axes (b, b1).
When the wings are raised
in the upper position (c, c1),
they are rapidly returned in
the start position (a, a1)

J Insect Behav

copula, the male and the female were largely immobile, with periodical male body
rocking and legs scrubs. After mating, both sexes frequently displayed a body cleaning
behavior. Non receptive females did not remain still and avoided the males jump. B.
oleae males did not show any preference for virgin or mated females (Fig. 3).
Wing Vibration Video Characterization
Male wing vibration parameters in successful and unsuccessful matings are reported
in Table 1. Frequency (t07.199, P<0.0001), pulse duration (t04.402, P<0.0001), the
distance between the two flies before the males jump (t05.056, P<0.0001), and the
female immobility (t06.368, P<0.0001) were higher in successful mating with
respect to unsuccessful ones. In successful mating attempts, male angles measured
during courtship and before the males jump were significantly different from those
performed in unsuccessful ones (t01.858, P00.0347 and t07.157, P<0.0001, respectively). As concerns wing sexual dimorphism, male wings were significantly
longer (t04.913, P<0.0001) and the wing posterior portion area was broader in males
with respect to females (t010.625, P<0.0001) (Table 2; Fig. 4).
Role of Male Wing Vibration in Mating
When males wings were removed the percentage of successful mating was significantly
reduced both in presence of virgin and mated females (2 025.62, P<0.001 and
2 021.68, P<0.001, respectively) (Fig. 5).

Discussion
Our results showed that the B. oleae mating behavior was very similar to Bactrocera
dorsalis (Poramarcom 1988; Poramarcom and Boake 1991). In fact, in both species
three main phases can be identified: attraction, courtship, and copulation. With regard

Fig. 3 The number of B. oleae males (virgin or mated) that displayed wing vibration (a) and copulation
attempts (b), in presence of virgin or mated females (n050; GLM, 2 post hoc test, P<0.05)

DistCA
(mm)

Mean values (s) are followed by the relative standard deviation (n030; Students t-test, P<0.05)

Frequency
(Hz)

Inter pulse
interval (s)
0.230.13 a 0.360.09 a

Pulse
duration (s)

31.810.88 a 320.8912.95 b 0.120.05 b 0.380.08 a

31.880.80 a 340.717.72 a

immobility Amplitude
(s)
()

9.811.73 a 7.221.23 a 5.540.87 a

DistWV
(mm)

12.674.23 b 76.1818.94 a 12.613.47 b 43.9423.10 a 10.052.22 a 5.511.38 b 3.371.65 b

CA ()

UM

CA ()

15.507.20 a 77.2421.24 a 20.775.19 a 50.9612.65 a

WV ()

SM

Trial WV ()

Table 1 Variables measured in B. oleae male wing vibrations (WV) before a successful (SM) or unsuccessful mount (UM). WV : the angle of male orientation toward the
female during wing vibrations; WV : the angle of alignment between male and female during wing vibrations; CA : the angle of male orientation toward the female
immediately prior to copulation attempt; CA : the angle of alignment between male and female immediately prior to male copulation attempt; distWV: distance between the two
flies (prothorax-to-prothorax) during wing vibrations; distCA: distance between the two flies prothorax prior to male jump. SM: successful; UM: unsuccessful mount

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Table 2 Parameters measured in B. oleae wings of both sexes
Wing Length (m)

Width (m)

Total area (m2)

Posterior portion
area (m2)

[Posterior portion
area/Total area]
*100

5176.27180.46 a 2491.1794.37 a 7738.22685.33 a 3976.58405.46 a 51.463.00 a

4930.6314.93 b

2464.2284.60 a 7016.51443.01 b 3087.16219.57 b 43.981.25 b

Mean values (s) are followed by the relative standard deviation (n050; Studentst-test, P<0.05)

to the C. capitata mating behavior, the olive fruit fly courtship differed in terms of the
complete absence of male wing buzzing (Briceo et al. 1996), head rocking (Webb et
al. 1983; Briceo et al. 1996), and aristae tapping before the copulation attempt
(Miranda 2000). It was observed that the female mating status did not affect male
preferences. This may be correlated with the female production of sex attractants that
does not cease or diminish after mating (Canale et al. 2010).

Fig. 4 A schematic representation of B. oleae male and female wings. Wing length: the distance from the
proximal margin of the alula (a) to the distal tip of the wing (b). Width: distance from the anterior edge of
the wing (c) to the posterior most extension of the anal lobe (d). The circle dotted line surrounded the
posterior portion of the wing

J Insect Behav

Fig. 5 Percentage of successful mating displayed by B. oleae virgin males, (wingless or not) in presence of
virgin (a) or mated (b) females. (n060; GLM, 2 post hoc test, P<0.05)

Our study provided behavioral evidence of the importance of male wing vibration
during courtship for successful mating. Firstly, when the males did not make a wingvibration courtship, but simply jumped onto the female and immediately attempted to
copulate, no successful matings were registered. Secondly, when the males wings
were removed, the successful matings were significantly reduced. This is in agreement with the findings observed in C. capitata by Keiser et al. (1973) and in the
Drosophila genus (Ewing 1964; Wilkinson 1987; Hoikkala and Aspi 1993). In
contrast, in other Diptera species wing-amputated males can produce an almost
normal courtship song (Waldron 1964; Miller et al. 1975; Tomaru and Oguma
1994). In Drosophila silvestris and D. heteroneura it was noted that wing ablation
did not affect male mating success at all (Boake and Poulsen 1997). Thirdly, there
were significant differences in frequency and pulse duration of the males wing
vibration during courtship in successful matings compared to unsuccessful ones.
According to findings on A. suspensa (Webb et al. 1984), wing vibrations performed
by successful males have a higher frequency than unsuccessful ones. In contrast, the
observed positive correlation between longer pulse duration and success in mating
disagrees with the findings observed in Drosophila littoralis and D. montana (Aspi
and Hoikkala 1995). Finally, the finding that the B. oleae females immobility before
the male jump was longer in successful matings is at variance with what has been
found in C. capitata (Briceo and Eberhard 2002). Data on the critical role of wing
vibration during courtship are in agreement with much other evidence. In C. capitata,
it was observed that mounts by wild males that failed in copulation were preceded by
significantly shorter courtships (Briceo and Eberhard 2000a). In B. dorsalis, females
were less attracted to those males that spent less time performing wing vibrations, and
the males ability to produce continuous wing vibration signals over a long time was
important in determining success in mating (Poramarcom 1988; Poramarcom and
Boake 1991).
It is important to note that in several Diptera species, wing vibration
parameters, such as frequency and amplitude, were related to male size. In A.
suspensa, calling propensity was significantly higher and calling song pulse train
interval was significantly shorter in large males (Burk and Webb 1983). Similarly, in
D. melanogaster it was noted that larger males had faster mating speeds than smaller
males (Partridge and Farqhuar 1983). In contrast, in other Diptera no significant
relation between male body size and mating success was found (Aspi and Hoikkala

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1995; Briceo and Eberhard 2002). In Drosophila planitibia very low correlations
were found between wing length and courtship song frequency, as well as between
wing length and burst length (Hoikkala and Kaneshiro 1997). However, we cannot
rule out that male body size - as well as several other factors such as age (Miyatake
and Kanmiya 2004), diet (Diamantidis et al. 2008), geographical origin (Briceo et al.
2002), and mass-rearing (Calcagno et al. 1999) - can affect wing vibration parameters
and success in mating in the olive fruit fly. Further research is required to clarify this
point.
Concerning wing sexual dimorphism, our data has shown that B. oleae male wings
are broader in males than in females, in agreement with a previous study on C.
capitata (Briceo and Eberhard 2000b). According to the hypothesis formulated by
Briceo and Eberhard (2000b) for medfly, also in B. oleae wing sexual dimorphism
could reflect an adaptation for physical cue production. On the other hand, the
enlarged posterior portion of the males wing may enhance the release of volatile
substances from the urotergal glands during wing vibration (Raspi et al. 1997). In B.
dorsalis, Poramarcom (1988) has demonstrated that during courtship the male wing
vibration transmits acoustic and olfactory cues that are both necessary for attracting
females.
We believe that the results of this work increase knowledge of the sexual behavior
of olive fruit fly and explain the functional role of male wing vibration. Further
research is required to acoustically characterize these male-borne cues.
Acknowledgements We are grateful to two anonymous reviewers for their useful comments on the
manuscript, Dr. Adrian John Wallwork for proofreading the English, Dr. Francesco Lanzo for providing the
drawing of wing movements, Dr. Augusto Loni and Paolo Giannotti for technical assistance.

References
Aspi J, Hoikkala A (1995) Male mating success and survival in the field with respect to size and
courtship song characters in Drosophila littoralis and D. montana (Diptera: Drosophilidae). J Ins
Behav 8:6787
Baker R, Herbert RH, Howse PE, Jones OT, Francke W, Reith W (1980) Identification and synthesis of the
major sex pheromone of the olive fly, Dacus oleae. J Chem Soc 1:5254
Boake CRB, Poulsen T (1997) Correlates versus predictors of courtship success: courtship song in
Drosophila silvestris and D. heteroneura. Anim Behav 54:699704
Briceo RD, Eberhard WG (2000a) Possible Fisherian changes in female mate-choice criteria in a massreared strain of Ceratitis capitata (Diptera: Tephritidae). Ann Entomol Soc Am 93:343345
Briceo RD, Eberhard W (2000b) Male wing positions during courtship by Mediterranean fruit flies
(Ceratitis capitata) (Diptera: Tephritidae). J Kans Ent Soc 73:111115
Briceo RD, Eberhard W (2002) Decisions during courtship by male and female medflies (Diptera:
Tephritidae): correlated changes in male behavior and female acceptance criteria in mass reared flies.
Fla Entomol 85:1431
Briceo RD, Ramos D, Eberhard W (1996) Courtship behaviour of male Ceratitis capitata (Diptera:
Tephritidae) in captivity. Fla Entomol 79:130143
Briceo RD, Eberhard WG, Vilardi JC, Liedo P, Shelly TE (2002) Variation in the intermittent buzzing
songs of male medflies (Diptera: Tephritidae) associated with geography, mass-rearing, and courtship
success. Fla Entomol 85:3240
Briceo RD, Eberhard W, Vilardi J, Cayol JP, Shelly T (2007) Courtship behaviour of different wild strains
of Ceratitis capitata (Diptera: Tephritidae). Fla Entomol 90:1518
Briscoe AD, Chittka L (2001) The evolution of color vision in insects. Annu Rev Entomol 46:471510

J Insect Behav
Burk T (1981) Signaling and sex in acalyptrate flies. Fla Entomol 64:3043
Burk T, Webb JC (1983) Effect of male size on calling propensity, song parameters and mating success in
Caribbean fruit flies, Anastrepha suspensa (Loew) (Diptera: Tephritidae). Ann Entomol Soc Am
76:678682
Calcagno GE, Vera MT, Manso F, Lux SA, Norry FM, Munyiri FN, Vilardi JC (1999) Courtship behaviour
of wild and mass-reared Mediterranean fruit fly (Diptera: Tephritidae) males from Argentina. J Econ
Entomol 92:373379
Canale A, Carpita A, Conti B, Canovai R, Raspi A (2010) Effect of age on 1,7-dioxaspiro-[5.5]-undecane
production in both sexes of olive fruit fly, Bactrocera oleae (Rossi) (Diptera Tephritidae). In:
Proceedings of the meeting Integrated Protection of Fruit Crops, Vico del Gargano, September
1217 2010- IOBC wprs Bulletin, Bulletin OILB srop 72:219225
Cavalloro R, Delrio G (1970) Rilievi sul comportamento sessuale di Dacus oleae (Gmelin) (Diptera,
Tripetidae) in laboratorio. Redia LII: 201230
Colegrave N, Hollocher H, Hinton K, Ritchie MG (2000) The courtship song of African Drosophila
melanogaster. J Evol Biol 13:143150
Daane KM, Johnson MW (2010) Olive fruit fly: managing an ancient pest in modern times. Annu Rev
Entomol 55:151169
De Marzo L, Nuzzaci L, Solinas M (1978) Studio anatomico, istologico, ultrastrutturale e fisiologico del
retto ed osservazioni etologiche in relazione alla possibile produzione di feromoni sessuali nel maschio
di Dacus oleae (Gmelin). Entomologica (Bari) XIV: 203266
Diamantidis AD, Papadopoulos NT, Carey JR (2008) Medfly populations differ in diel and age patterns of
sexual signaling. Ent Exp Appl 128:389397
Dukas R, Mooers A (2003) Environmental enrichment improves mating success in fruit flies. Anim
Behav 66:741749
Ewing AW (1964) The influence of wing area on the courtship behaviour of Drosophila melanogaster.
Anim Behav 12:316320
Feron M, Andriew AJ (1962) Etude des signaux acoustiques du male dans le comportement sexual des
Dacus oleae Gmel. (Diptera: Trypetidae). Ann Epiphyt 13:269276
Gen H, Nation JL (2008) Maintaining Bactrocera oleae (Gmelin.) (Diptera: Tephritidae) colony on its
natural host in the laboratory. J Pest Sci 81:167174
Haniotakis GE (1974) Sexual attraction in the olive fruit fly Dacus oleae (Gmelin). Environ Entomol 3:82
86
Haniotakis GE (1977) Male olive fly attraction to virgin females in the field. Ann Zool-Ecol Anim 9:273276
Hoikkala A, Aspi J (1993) Criteria of female mate choice in Drosophila littoralis, D. montana, and D.
ezoana. Evolution 47:768777
Hoikkala A, Kaneshiro KY (1997) Variation in male wing song characters in Drosophila planitibia
(Hawaiian picture-winged Drosophila group). Anim Behav 10:425436
Joyce AL, Aluja M, Sivinski J, Vinson SB, Ramirez-Romero R, Bernal JS, Guillen L (2010) Effect of
continuous rearing on courtship acoustics of five braconid parasitoids, candidates for augmentative
biological control of Anastrepha species. BioControl 55:573582
Keiser I, Kobayashi RM, Chambers DL, Schneider EL (1973) Relation of sexual dimorphism in the wings,
potential stridulation, and illumination to mating of oriental fruit flies, melon flies, and Mediterranean
fruit flies in Hawaii. Ann Ent Soc Am 66:937941
Kyriacou CP, Hall JC (1982) The function of courtship song rhythms in Drosophila. Anim Behav 30:794
801
Loher W, Zervas G (1979) The mating rhythm of the olive fruit fly Dacus oleae (Gmelin). Z ang Ent
88:425443
Mavraganis VG, Papadopoulos NT, Kouloussis NA (2010) Extract of olive fruit fly males (Diptera:
Tephritidae) attract virgin females. Entomol Hell 19:1420
Mazomenos BE (1989) Dacus oleae. In: Helle W (Ed) Word crop pests Vol. 3A: fruit flies: their biology,
natural enemies and control. p 169
Miller DD, Goldstein RB, Patty RA (1975) Semispecies of Drosophila athabasca distinguishable by male
courtship sounds. Evolution 29:531544
Miranda X (2000) Sexual dimorphism in the aristae of Ceratitis capitata (Diptera, Tephritidae) and its
possible importance in courtship. J New York Entomol Soc 108:339348
Miyatake T, Kanmiya K (2004) Male courtship song in circadian rhythm mutants of Bactrocera cucurbitae
(Tephritidae: Diptera). J Ins Physiol 50:8591
Partridge L, Farqhuar M (1983) Lifetime mating success of female fruit flies (Drosophila melanogaster) is
related to their size. Anim Behav 31:871877

J Insect Behav
Poramarcom R (1988) Sexual communication in the Oriental fruit fly, Dacus dorsalis Hendel (Diptera:
Tephritidae). Ph.D. Thesis, University of Hawaii, Honolulu
Poramarcom R, Boake CRB (1991) Behavioural influences on male mating success in the Oriental fruit fly,
Dacus dorsalis Hendel. Anim Behav 42:453460
Raspi A, Canale A, Lucchi A (1997) Preliminary observations on integumentary urotergal glands in
Bactrocera oleae (Gmelin) (Diptera Tephritidae). Frustula entomol n.s. XX (XXXIII):193202
Rice RE, Phillips PA, Stewart-Leslie J, Sibbett GS (2003) Olive fruit fly populations measured in central
and southern California. Calif Agric 57:122127
Sivinski J, Burk T, Webb JC (1984) Acoustic courtship signals in the Caribbean fruit fly, Anastrepha
suspensa (Loew). Anim Behav 32:10111016
Sivinski J, Calkins CO, Webb JC (1989) Comparison of acoustic courtship signals in wild and laboratory
reared Mediterranean Fruit Fly Ceratitis capitata. Fla Entomol 72:212214
Sprinthall RC (2003) Basic statistical analysis. Pearson Education, New Jersey
Tomaru M, Oguma Y (1994) Differences in courtship song in the species of the Drosophila auraria
complex. Anim Behav 47:133140
Tzanakakis ME (2006) Insects and mites feeding on olive: distribution, importance, habits, seasonal
development and dormancy. Brill Acad Publ Leiden
Tzanakakis ME, Tsitsipis JA, Economopoulos AP (1968) Frequency of mating in females of the olive fruit
fly under laboratory conditions. J Econ Entomol 61:13091312
von Rolli K (1976) Die akustischen Sexualsignale von Ceratitis capitata Wied. und Dacus oleae Gmel. Z
ang Ent 81:219223
von Schilcher F (1976) The function of pulse song and sine song in the courtship of Drosophila
melanogaster. Anim Behav 24:622625
Waldron I (1964) Courtship sounds production in two sympatric sibling Drosophila species. Science
144:191193
Webb JC, Sharp L, Chambers DL, McDow JJ, Benner JC (1976) Analysis and identification of sounds
produced by the male Caribbean fruit fly, Anastrepha suspensa. Ann Entomol Soc Am 69:415420
Webb JC, Calkins CO, Chambers DL, Schwienbacher W, Russ K (1983) Acoustical aspects of behaviour of
Mediterranean fruit flies. Ceratitis capitata: analysis and identification of courtship sounds. Entomol
Exp Appl 33:18
Webb JC, Sivinski J, Litzkow C (1984) Acoustical behaviour and sexual success in the Caribbean Fruit Fly,
Anastrepha suspensa (Loew) (Diptera: Tephritidae). Environ Entomol 13:651656
Wilkinson GS (1987) Equilibrium analysis of sexual selection in Drosophila melanogaster. Evol 41:1121
Zervas GA (1982) Reproductive physiology of Dacus oleae (Gmel.) (Diptera: Trypetidae). Comparison of
a wild and artificially reared flies. Geoponica (in Greek) 282:1014
Zouros E, Krimbas CB (1970) Frequency of male bigamy in natural population of the olive fruit fly Dacus
oleae as found by using enzyme polymorphism. Entomol Exp Appl 13:19