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Analysis of the Territorial, Courtship and Coupling Behavior of the Hemipterophagous Butterfly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae)


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In this study we examined the territorial, courtship and coupling behavior, mating interruptions and influence of sunlight on mating behavior in Spalgis epius (Westwood). We constructed an ethogram of territorial and courtship behavior, and catalogued the sequence of behavioral acts associated with mating behavior. Mating behavioral acts were divided into four repertoires i.e. pre-courtship, pre-coupling, coupling and post-coupling behaviors. Totally 22 behavioral acts were recorded from four repertoires. Tree canopy, canopy height and sunlight conditions are the important factors that influence copulation in S. epius. The courtship activity led to successful copulation in 71.9 % pairs. Incidence of different types of courtship and copula interruptions in S. epius was also recorded.
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Analysis of the Territorial, Courtship and Coupling
Behavior of the Hemipterophagous Butterfly, Spalgis
epius (Westwood) (Lepidoptera: Lycaenidae)
Anegunda S. Dinesh &Melally G. Venkatesha
Revised: 7 May 2012 /Accepted: 21 May 2012 /
Published online: 31 May 2012
#Springer Science+Business Media, LLC 2012
Abstract In this study we examined the territorial, courtship and coupling behavior,
mating interruptions and influence of sunlight on mating behavior in Spalgis epius
(Westwood). We constructed an ethogram of territorial and courtship behavior, and
catalogued the sequence of behavioral acts associated with mating behavior. Mating
behavioral acts were divided into four repertoires i.e. pre-courtship, pre-coupling,
coupling and post-coupling behaviors. Totally 22 behavioral acts were recorded from
four repertoires. Tree canopy, canopy height and sunlight conditions are the important
factors that influence copulation in S. epius. The courtship activity led to successful
copulation in 71.9 % pairs. Incidence of different types of courtship and copula
interruptions in S. epius was also recorded.
Keywords Spalgis epius .territory .perching .courtship .mating .mealybugs
Different butterfly species exhibit considerable diversity in the pattern of their mating
systems (Gilbert 1976; Wiklund 1977; Davies 1978). Generally males initiate sexual
behavior in several butterfly species and visually locate females (Magnus 1958;
Obara 1970; Hidaka and Yamashita 1975). Male butterflies adopt four main mate
locating strategies i.e. perching, patrolling, territorial defense, and lek assembly
(Shields 1967; Baker 1984; Scott 1974; Davies 1978; Rutowski 1991). Very little is
known about mate location sites and behavior in butterflies (Rutowski 1991). In
J Insect Behav (2013) 26:149164
DOI 10.1007/s10905-012-9341-9
A. S. Dinesh :M. G. Venkatesha (*)
Insect Science Laboratory, Department of Zoology, Bangalore University, Bengaluru 560 056, India
perching butterflies initial approach is mainly based on visual stimuli particularly on
female movement and size, which is helpful to discriminate between same and
different species (Tinbergen 1941). Once a male butterfly locates a receptive conspe-
cific female, aerial or ground based courting behavior will follow depending on
species (Pinzari 2009). Investigations on mating behavior in many butterfly species
of different families including Lycaenidae have revealed that territorial behavior is
primarily a perching activity (Alcock 1983; Cordero and Soberon 1990; Fischer and
Fiedler 2001). A variety of courtship behavior has evolved in response to mate choice
by female butterflies to choose those males that show specific characteristics, which
enhance the reproductive success of females (Pinzari 2009). Studies on mating
behavior and copulation mechanisms are helpful to understand the evolution of
mating systems (Thornhill and Alcock 1983) and in Lepidoptera it is useful to
examine how ecological conditions curtail or promote sexual selection and the
evolution of different mating systems (Hughes et al. 2000). Moreover, to exploit
important entomophagous insects as biocontrol agents, knowledge of their mating
behavior is essential to plan an appropriate mass rearing program (van den Assem and
Povel 1973).
The apefly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae: Miletinae) has
been recorded as a predator of various species of mealybugs viz., Planococcus
virgatus (Cockerell), P. lilacinus (Cockerell), P. citri (Risso), Ferrisia virgata (Cock-
erell), Maconellicoccus hirsutus (Green) and Paracoccus marginatus Williams and
Granara de Willink (Hemiptera: Pseudococcidae); and scale insects Chloropulvinaria
polygonata (Cockerell.) (Hemiptera: Coccidae) and Dactylopius sp. (Hemiptera:
Dactylopiidae) (see Dinesh and Venkatesha 2011a). Moreover, S. epius has been
reported as a potential predator of P. citri, P. lilacinus and M. hirsutus in India
(Chacko et al. 1977; Gowda et al. 1996; Rahiman and Vijayalakshmi 1998; Venkatesha
and Shashikumar 2006). Dinesh and Venkatesha (2011a,b) have reported that S.
epius can be used as a major biocontrol agent of M. hirsutus and P. citri.
Spalgis epius occurs in India, Bangladesh, Burma, Sri Lanka, Philippines, Java,
Thailand and Krakatau Island (Indonesia) (see Dinesh and Venkatesha 2011a). The
morphology, life history and feeding potential of S. epius has been studied (De
Niceville 1890; Aitken 1894;Bingham1907; Vinod Kumar et al. 2006,2008a;
Dinesh et al. 2010; Thangamalar et al. 2010; Dinesh and Venkatesha 2011a,b).
Observations on the activities of S. epius and its interaction with ants have been made
in the field (Venkatesha et al. 2004; Venkatesha 2005; Vinod Kumar et al. 2008b).
Venkatesha and Dinesh (2011) developed the first mass rearing method for S. epius.
Very little is known about the courtship and mating behavior of S. epius (Dinesh et
al. 2010; Venkatesha and Dinesh 2011). Moreover, no information on the mating
behavior of any other carnivorous lycaenid species is available. Hence, we made
observations on the territorial, courtship and mating activities of S. epius both in
captivity and field. In this paper we present a detailed ethogram of mating behavior
including courtship and copula interruptions and influence of sunlight on mating in S.
epius. This study provides the first clear knowledge about the mating behavior of a
hemipterophagous butterfly species and it is useful for investigations on the biology
and rearing of other economically important carnivorous lycaenid species. Besides,
this study also helps to broaden our knowledge about mating behavioral patterns in
phytophagous and carnivorous lycaenids.
150 J Insect Behav (2013) 26:149164
Materials and Methods
The study was carried out in the Bangalore University campus, Bengaluru, India
(Latitude 12° 58N, longitude 77° 35E, elevation 921 masl).
Lab Rearing of S. epius
To rear S. epius in the laboratory, its prey P. c i t r i was cultured on pumpkins
(Cucurbita maxima Duchesne) following the method of Serrano and Lapointe
(2002). Spalgis epius was reared on the mealybug infested pumpkins at 28 ± 1°C
and 65±5 % relative humidity as described by Dinesh et al. (2010). Spalgis epius
adults reared in the laboratory were utilized for conducting studies on courtship
behavior. Freshly eclosed adults were fed with water in the laboratory to enhance
their longevity (Dinesh et al. 2010).
The Ethogram of Mating Behavior
To make an ethogram of mating behavior of S. epius, observations were made in an
outdoor nylon tent cage (6 m length×6 m width× 10 m height) placed over a tree
(9 m height) in the field as followed by Venkatesha and Dinesh (2011). As S. epius
adults are known to mate a day after their pupal eclosion (Dinesh et al. 2010), for
each observation five to ten pairs of 1 day old S. epius adults reared in the laboratory
were released into the cage at 0700 h and their territorial, courtship and mating
behavior was observed during their active period (i.e. 08001700 h) till their death.
Based on the focal-animal observation method (Altmann 1974) behavioral acts and
duration of each act of males and females directly involved in mating were recorded
and described the sequences of acts. The sequence of mating behavior was described
using all-occurrences sampling method (Altmann 1974). Altogether randomly select-
ed 135 pairs were released into the cage for observations on courtship behavior. Close
observations on every courtship event in the cage was made from a plank platform
(6 m height), which was set up adjacent to the tree canopy as followed by Venkatesha
and Dinesh (2011). In addition to visual observations a video camera (Sony HDR-
XR150E, 25× optical zoom lens) was set up to record all details of focal-animal
courtship activities. Later, video and visual recordings were critically analyzed and
noted every details of courtship behavior. Duration of territorial and courtship flights
was recorded using a stop watch (Model-EC8071). To differentiate between male and
female butterflies during courtship observations, ventral side of both forewings of
males and females were marked with black and red marker ink (Reynolds, India ink),
respectively, before releasing them into the mating cage. Moreover, males and
females of S. epius can be differentiated based on shape of forewings (Dinesh et al.
2010). To distinguish different territorial males of S. epius during courtship observa-
tions, forewings of males were marked with black marker ink at selected spots.
Identity of resident or intruder males in a territory was confirmed by using a
binocular (Nikon 12x50) or zoom in picture of video recording based on the
position of ink markings on the forewing. Daytime ambient temperature within
the tree canopy in the cage was recorded by a digital thermometer (Temp.Tec:
J Insect Behav (2013) 26:149164 151
Mating Interruptions
The occurrence of courtship and copula interruptions in captivity was recorded.
Courtship Behavior in the Field
Five to ten pairs of 1 day old S. epius adults reared in the laboratory were released
each observation period on bushes/trees in the field and recorded their courtship
behavior. Totally 50 pairs were released in the field. In addition, observations were
made on courtship behavior of the field population of S. epius in a mealybug (P.
marginatus) infested papaya (Carica papaya L.) field in Bengaluru, where a high
incidence of S. epius was noticed.
Influence of Time and Daylight on Mating
To assess the peak mating time and successful mating under different daylight
conditions i.e., bright/diffused sunlight and overcast sky, five pairs of 1 day old S.
epius were released into the cage for 5 days under each of the above conditions. The
copulation time and number of pairs copulated under different daylight conditions
were recorded.
Data Analysis
To test the difference in the number of pairs copulating at different timings of the day
under different daylight conditions, one-way ANOVA was conducted and significant
difference was determined utilizing Tukey HSD test at P5 % (SPSS Inc. 2008).
The Ethogram of Mating Behavior in Captivity
Male and female butterflies of S. epius were active 1 day after their pupal eclosion.
Behavioral acts associated with mating behavior in S. epius were divided into four
repertoires i.e. pre-courtship, pre-coupling, coupling and post-coupling behaviors.
The number of behavioral acts recorded under each repertoire was 6, 6, 5 and 5,
respectively (Table 1). Of the total 22 behavioral acts, 13 and two were performed
exclusively of male and female, respectively, and seven acts common to both sexes.
The male initiated entire process of mating through perching and marking in the tree
canopy at 69 m (Mean±SE: 7.5 ± 0.9) height from the ground and established a
territory (Fig. 1). A single territorial male marked 11.2±3.4 perch points in the
territory; distance between perch points was 2025 cm. Spalgis epius males per-
formed territorial flights between 1100 and 1400 h. Males exhibited intense pre-
courtship flight activity in the canopy interspersed with short periods of basking.
Males spent most of their time in establishing territory, chasing intruder males and
courting females. Sometimes, a few S. epius males established territories simulta-
neously and strictly maintained a neutral space of 0.52 m between territories. In an
152 J Insect Behav (2013) 26:149164
Table 1 Description of ethogram for behavioral acts involved in the territorial, courtship and mating of
Spalgis epius
Behavioral act Sex Description
Territory marking Male The male marks a territory in a tree canopy by dragging his
abdominal tip on the foliages of proximal end of the twigs.
Perching Male The male perches at some fixed points in a horizontal plane
within the radius of one meter (Fig. 1).
Inspection Male The male occasionally makes spontaneous short flights in a limited
range within the 23 m radius around the perch points in absence
of any conspecific individuals and frequently changes perch spots
within the marked territorial area.
Basking Male During perching activity the territorial male basks for
23 min by horizontally spreading both wings (Fig. 2a).
Chasing an intruder
by straight flight
Male Short flight wherein the territorial male chases a conspecific
intruder male in the territory in a horizontal plane without
any contest (Fig. 1).
Chasing an intruder
by circling flight
Male Long flight wherein the territorial male rushes towards a
conspecific intruder male in the territory and engages in an
ascending circling flight up to 11.5 m height from the
territory with 4-6 repeat bouts (Fig. 1).
Territorial flight Female The flying female enters the territory or the female sitting
in the territory starts flying (Fig. 2b).
Flight pursuit Male The territorial male engages in courtship flight almost immediately
after a flying female enters the territory (Fig. 2c).
Initiation of courtship
Male The pair initiates courtship flight in a horizontal plane
above the tree branches and then fly through the canopy.
Courtship flight Male During the courtship flight the male follows closely at 1218 cm
behind, but slightly above the steadily flying female (Fig. 2d).
Tracking Male The male follows the female constantly without losing the track till
the end of courtship flights.
Alighting on vegetation Male First the female alights suddenly on the foliage and soon after the
male alights at a distance of 48 cm behind the female (Fig. 2e).
Crawling Male The alighted male walks alongside the stationary female.
Abdomen raising Female The alighted female raises her abdomen.
Parallel position Male The male passes alongside until the couple in close parallel head to
head position (Fig. 2f).
Abdominal bending Male The male quickly bends his abdominal tip to establish contact with
the genitalia of the female (Fig. 2g).
Coupling Male The male pushes its bent abdominal tip against the raised
female genitalia and interlocks.
Couple rotation Male After genital coupling the male moves laterally until both sexes face
opposite direction.
Couple abdomen raising Male Abdomens of the pair are held in slightly raised position
with their wings dorsally folded and heads pointing
downward all through the copulation (Fig. 2h).
J Insect Behav (2013) 26:149164 153
ascending circular territorial flight between residents and conspecific intruder males
(N015), resident males won and intruders were chased away. But, in short flight an
intruder male was chased away without contest (Table 1).
The sequence of successful courtship activities observed in S. epius is pre-
sented in Fig. 2. An active male attracted a virgin female within 23.8±7.8 min after
establishing a territory. The pair alighted after 14 rounds of courtship flights at an
interval of 45 min; each courtship flight last for about a minute. Sometimes pairs
made one or more alighting attempts before final settling. During courtship flight
males were very persistent and ceasing their activity only after females alighted.
Various courtship events and their duration in S. epius are presented in Fig. 3.
Duration of overall flight activity (i.e. perching and territorial activity, chasing an
intruder male, and courtship flight) of males was longer than that of females (see
Fig. 3).
In the final stage of courtship (N0104), 93.3 % (097) of S. epius males that
alighted behind females crawled to the left side of the females, then bent their
Table 1 (continued)
Behavioral act Sex Description
Pair in copula Male The coupled pair stays motionless and copulation lasts for
55.7± 9.6 min.
Flight Male Only in the instance of any disturbances by predators (i.e.
ants or spiders)/strong wind the pair flies in copula to different
place in the canopy. If it rains the mating pair moves underside
the leaf without interrupting the copulation (n06).
Copula termination Female Copulation terminated by slight movement of the female, thus
copula unplugs. Generally the separated pair remains in the
mating place for 1520 min. If it is an ending stage of the days
activity then they stay in the mating place for 3045 min.
Fig. 1 Territorial activity of Spalgis epius male butterfly. Dark grey: medium to thick canopy, Light grey:
thin canopy, White: space in the canopy, Dashed circle: territorial area, Crosses: perch points in the
territory. Straight line: straight flight in chasing of an intruder, Curved line: ascending circling flights
between resident and intruder males
154 J Insect Behav (2013) 26:149164
abdominal tip to the right and established contact with the female genitalia (Fig. 2f,
g). Only seven out of 104 males moved to the right of the females and in two of these
cases the males had trouble in establishing genital contact and the female ended flying
away. However, one of the two males in the second attempt quickly alighted on the
left side of the female and able to mate by bending its abdominal tip to its right side.
After genital coupling the male turns laterally until the pair reaches the tail-to-tail
position with slightly raised abdomens and dorsally folded wings (Fig. 2h). Coupling
is the longest phase of mating activity (see Fig. 3) and is characterized by the great
immobility of the couple at the mating site. Copulations constantly took place in the
canopy in the diffused sun shining area. Always copulation terminated by slight
movement of females. Totally 71.9 % (N0135) of pairs successfully copulated in
All the time S. epius pairs initiated courtship strictly inside the territory (N0104),
but 36.5 % of copulations took place within an own territory, 46.2 % in a neutral
space between territories, 5.8 % in a general area and 11.5 % in others territory.
Whenever a courting pair landed directly in others territory and copulated immedi-
ately, the resident male never disturbed the copulating pair. But, in two instances after
the pair detached, a resident male tried to court the mated female, which was strictly
Fig. 2 Sequence of acts of successful courtship and mating in Spalgis epius.aperching male in the
territory, bresting female in the tree canopy, cflying female in the territory, dcourtship flight of a pair, e
courted pair alighting on the vegetation, fcrawling male in parallel position with the courted female, g
copula joining (dorsal view), and hpair in copula
J Insect Behav (2013) 26:149164 155
Courtship Interruptions
A total of 128 courtship interruptions were recorded in S. epius in captive mating.
Wherein mainly five types of interruptions were observed: 1) multiple males courting
a single female- 33.6 % mating interruptions were because of multiple males trying to
court a single female. When many males were active in the cage they marked their
territory in close proximity to each other within the confined area. Soon after a female
entered the territory she was courted by the territorial male. Whenever the courting
pair got into others territory and continued their courtship flight, the resident male
suddenly followed the courting pair and when the courting female alighted on the
foliage, two competing males attempted to copulate the female, which flew away
under such circumstances. Any time a single female was courted by more than one
male, the courtship led to unsuccessful copulation. Although at times multiple males
courted a single female, 3.9 % of males that courted continuously without losing the
track of the female till the end led to successful copulation; 2) the male fails to locate
the female- 28.1 % of males courted the females till the end of the courtship flight and
finally when the receptive female quickly alighted on the foliage the courting male
was unable to locate the female; 3) losing the track of females- 18.0 % of courting
males lost the track of erratically flying females during courtship flight in the tree
canopy; 4) strong wind- strong wind disturbed the courtship flights and 7.0 % of
couples were unsuccessful in mating; and 5) obstacle- 13.3 % of courtship flights
Fig. 3 Courtship and mating behaviors showing various events and their duration in Spalgis epius. tt: tail
to tail
156 J Insect Behav (2013) 26:149164
were interrupted when courting couples had hit the wall of the mating cage and lost
courtship momentum.
Copula Interruption
After the successful union of 103 pairs of S. epius, only 5.8 % (06) of copula
interruptions were observed. In which 2.9 % pairs detached from their copula within
two minutes after copulation. In 1.9 % of courtship flights, both male and female
butterflies alighted on a leaf and then the male with difficulty made contact with the
unnaturally raised female genitalia, but the female detached from him and flew away.
In 1.0 % of copulations, 12 min after union the female began dragging the male
behind her and even she made small flights while dragging him. When the male was
still motionless, the female began twisting her body while dragging and in the process
the male was hanging in copula, and finally the pair got detached.
Courtship and Mating Behavior in the Field
When S. epius pairs were released in the field, they flew short distances and sat on the
tree foliages. Generally we could not track all the field released S. epius adults as they
flew away and lost in the sylvan. Hence, courtship and mating was observed only in
six pairs of the 50 pairs released on different days in the field. Although courting pairs
of S. epius were observed many times in the papaya field, mating was observed only
in four pairs on different days. The territorial, courtship and mating behavior of the
field released S. epius adults or in the natural population of S. epius in the papaya
field was similar to that observed in the captivity. But, the territory height range was
615 m from the ground in the field. Moreover, the ascending circling flight between
resident and intruder males was 46 m height from the territory.
Influence of Time and Sunlight on Mating
Under bright sunlight, the maximum number of pairs copulated between 11.00 and
12.00 h, though it was not significantly different from that mated between 10.00 and
11.00 h (Tukey HSD test: P00.115). However, these were significantly higher than
those (F
3, 16
017.939; P<0.05) mated during 12.0013.00 and 13.0014.00 h
(Fig. 4). In diffused sunlight, the pairs never mated between 10.00 and 11.00 h.
But, more pairs mated between 12.00 and 13.00 h than between 11.00 and 12.00 and ,
13.00 and 14.00 h and it was significantly different (F
3, 16
021.030; P<0.05).
Similarly, the pairs never mated between 10.00 and 11.00 h under overcast sky.
Although the pairs mated in subsequent hours, a few pairs mated between 11.00 and
12.00 h and, it was significantly lower than the pairs mated between 12.00 and 13.00
and 13.00 and 14.00 h (F
3, 16
07.704; P<0.05). Overall, the maximum numbers of
pairs mated under bright sunlight, but it was not significantly different from that
mated under diffused sunlight. However, number of pairs mated under bright/diffused
light was significantly higher than that mated under overcast sky (F
2, 57
08.453; P<
0.05) (Fig. 5). Spalgis epius mated in the temperature range of 2634°C under bright/
diffused daylight in the cage. But, whenever temperature increased above 36°C in the
hot season, S. epius very rarely mated in the cage.
J Insect Behav (2013) 26:149164 157
Spalgis epius males performed more behavioral acts than females (see Table 1). In most
butterfly species it is the male who actively seeks mates, and females are less active
Fig. 4 Influence of sunlight on
Spalgis epius mating during dif-
ferent hours of the day. abright
sunlight, bdiffused sunlight, c
overcast sky. Values are mean±
SE; Bars with different letters
indicate the significant difference
between the hours within the
same daylight condition at P<
0.05 (one way ANOVA Tukey
HSD test)
Bright Diffused Overcast
ht condition
Mean number of copulation
Fig. 5 Influence of different sunlight conditions on Spalgis epius mating. Values are mean±SE; Bars with
different letters indicate the significant difference between different lighting condition at P<0.05 (one way
ANOVA Tukey HSD test)
158 J Insect Behav (2013) 26:149164
than males (Scott 1972; Rutowski 1984). The pre-courtship behavior of S. epius (i.e.
territorial marking and perching at fixed points) is similar to other phytophagous
lycaenids viz., Atlides halesus (Cramer) (Alcock 1983), Callophrys xami (Reakirt)
(Cordero 1993), Favonius taxila Bremer (Takeuchi and Imafuku 2005a), and Chrys-
ozephyrus smaragdinus Bremer (Takeuchi and Imafuku 2005b). The spontaneous
flight of S. epius territorial male in the territory may provide inspection of the territory
as reported in Lycaena hippothoe (Linnaeus) (Fischer and Fiedler 2001), F. taxila
(Takeuchi and Imafuku 2005a), and C. smaragdinus (Takeuchi and Imafuku 2005b).
More number of S. epius butterflies was able to mate in the cage. It may be because of
butterflies that adopt perching in the territories are known to have higher mating
success than patrolling ones (Wickman 1986).
Generally in butterfly species with more number of males than females perform
perching, territorial defense and lek assembly activities (Ehrlich 1984). However, in S.
epius though males perform perching and territorial defense, their number is always
lower than that of females (Vinod Kumar et al. 2006; Dinesh et al. 2010). Basking
behavior observed in S. epius territorial males during territorial flights is also reported
in other lycaenids e.g., Incisalia iroides (Boisduval) (Powell 1968), C. smaragdinus
(Takeuchi and Imafuku 2005b) and F. taxila (Takeuchi and Imafuku 2005a). Basking
behavior of territorial male butterflies is to regulate their body temperature as they
require high thoracic temperature in order to perform flight activities (Clench 1966;
Pivnick and Mcneil 1986). Moreover, Stutt and Willmer (1998) reported that resi-
dents can achieve a higher body temperature by basking in a territory, which enhances
their physical condition, and are therefore more likely to win a contest with an
intruder. Similar behavior was observed in S. epius. Likewise, resident males of
Pararge aegeria (Linnaeus) are almost always known to win (Davies 1978; Wickman
and Wiklund 1983). Takeuchi and Honda (2009) reported that real fighting ability is
correlated with ownership status. Furthermore, as butterflies are ectotherms, owners
win because they are able to rise their body temperature to optimum levels by basking
in a territory, whereas intruder males are not (Stutt and Willmer 1998). The two types
of chasing flights between territorial and intruder males of S. epius are also observed
in a lycaenid Eumaeus toxea Godart (Martinez-Lendech et al. 2007). The circling
ascending flight between resident and intruder males of S. epius is similar to that of A.
halesus (Alcock 1983), Strymon melinus Hübner (Alcock and ONeill 1986), C. xami
(Cordero 1993) and C. smaragdinus (Takeuchi and Imafuku 2005b). The circling
flights of territorial and intruder males of S. epius were initiated within a territory, and
invariably terminated outside the territory as observed with L. hippothoe (Fischer and
Fiedler 2001) and F. taxila (Takeuchi and Imafuku 2005a). Contest between individ-
ual males for mating opportunities is a widespread phenomenon and constitutes an
important factor of Darwins(1871) theory of sexual selection. Spalgis epius territo-
rial males maintained neutral spaces between territories as observed in various
butterfly species (Knapton 1985).
The initiation of courtship flight followed by approaching only a flying female by
a territorial male including entire courtship behavior in S. epius is similar to that of C.
xami (Cordero 1993)andC. smaragdinus (Takeuchi and Imafuku 2005b). This
indicates that visual signal plays an important role in S. epius courtship behavior as
observed in several species of butterflies (Lutz 2002; Jiggins et al. 2004; Oliver et al.
2009). Therefore, when a courting pair of S. epius gets into others territory the
J Insect Behav (2013) 26:149164 159
resident male joins the courting pair. But, if a courting pair lands directly in others
territory and copulate immediately, the resident male never disturbs the pair in copula.
However, we are not sure whether only visual signal or male pheromone or both are
involved in attraction of a female to the male territory in S. epius. The alighting and
copulating behavior of S. epius is similar to that of C. xami (Cordero 1993) and C.
smaragdinus (Takeuchi and Imafuku 2005b). The successful copulation of S. epius
males by bending abdominal tip to their right side is not reported so far in any other
species. Furthermore, it is not known why an S. epius male struggles to bend its
abdominal tip to the left when it rarely crawls to the right side of the female. As far as
we know our description of genital coupling behavior of males from the right or left
side of the female in S. epius is the first record of occurrence of this type of behavior
in any species of lycaenids in particular and other butterfly families in general. This
behavior seems to be crucial for the copulatory success of S. epius males because
when a male struggles to establish genital contact from the right side of the female,
she flies away.
As S. epius is a shy butterfly and hide in bushes (Venkatesha et al. 2004), its
courtship and mating activities are not commonly observed in the field. However, our
observations on the mating behavior of S. epius in the field have revealed that it is
almost similar to that in the confined condition. As we could observe courtship
activity in a few individual pairs of S. epius in the field, information on neutral
spaces including several other courtship events could not be recorded. The courtship
and mating activity of S. epius in the field may purely depends on the availability of
hemipteran host insects as the predator pupates on the hemipteran infested
bushes/trees and this short lived species (Dinesh et al. 2010) may mate in the
same place. However, if the host infestation in the field is on the very small bushes,
S. epius adults may fly to a different place to find a tree canopy as they mate at a
moderate height from the ground level. In phytophagous butterflies conditions like
the occurrence of larval host plants, temperature and heat, adult nectar resources, and
geography will increase the density of individuals and enhance encounter rates
between sexes (Dennis and Shreeve 1988). The same may be true in hemipteropha-
gous S. epius.
Several courtship interruptions in S. epius were because of multiple males courted
a single female in the cage. Whenever more numbers of S. epius pairs were present in
the cage the incidence of competition among males was more in a restricted space.
However, this condition was not observed in natural condition. When swift alighting
of a receptive S. epius female on the leaf coincided with a flying intruder male diverts
the concentration of the courting male that is unable to track the female. Strong wind
or some physical obstacles interrupted the courtship in S. epius as observed with C.
xami (Cordero 1993). Copulation of S. epius in the partly sun shining area in the
canopy may prevent harmful overheating or attack of visually oriented predators as
reported in the butterfly Dryas iulia alcionea F. (Mega and de Araujo 2010). In case
of disturbance from rain or predators, it is always females of S. epius in copula that
drag males to some other safe location.
Some pairs of S. epius separated within a few minutes after interlocking as
reported by Cordero (1993)inC. xami. Generally copula interruption occurs quickly
owing to rapid assessment of mate quality as physical separation may be difficult if
copulation is prolonged (Wickman 1985). Furthermore, copula interruptions in C.
160 J Insect Behav (2013) 26:149164
xami were due to mate choice after mating began (Cordero 1993). Females or males
may be able to evaluate their mating partners after copulation begins and decide to
interrupt it within a few minutes (Eberhard 1985,1994; Eberhard and Cordero 1995;
Cordero 1995).
The high incidence of mating under bright sunlight in butterflies is because of their
ability to absorb heat from the sunlight through their wings (Watt 1968) and the behavior
of ectothermic insects is constrained by ambient thermal conditions (Heinrich 1996).
Sunlight is crucial for flight behavior in butterflies since the performance of thoracic
flight muscle is dependent on the thoracic temperature, which in turn depends on the
ambient thermal conditions (Watt 1969; Kammer 1970; Rawlins 1980). Hence, under
bright/diffused sunlight S. epius males were active and marked the territory, under-
took territorial flights as well as chasing flights with intruder males and located active
females and had more copulations. Hence, the delayed mating activity in S. epius
under diffused and overcast conditions may be due to prolonged exposure to light to
rise their body temperature for courtship activities. Most pairs of S. epius mated under
bright/diffused sunlight than under overcast sky as observed in the lycaenid butterfly
Heodes virgaureae (L.) (Douwes 1976). However, S. epius pairs rarely mated under
rising temperature during summer period. This could be one of the reasons for low
population of S. epius during summer months as reported by Venkatesha and Sha-
shikumar (2006) and Vinod Kumar et al. (2006). Initiation of copulation separation
by S. epius females suggests that the duration of sperm transfer is determined by the
female as noticed in C. xami (Cordero 1993) and in some other species of butterflies
(Scott 1972) and several species of moths (Phelan and Baker 1990). However, in
several species of butterflies, it is males that determine the end of copulation
(Wickman 1985).
Much work is still to be done to explain the role of visual and chemical
signaling in courtship activity and male coupling behavior in S. epius. Yet, we
hope our study provides the first detail ethogram of the territorial, courtship and
mating behavior of a hemipterophagous butterfly. It is evident that under captivity S.
epius adults should be provided primarily a tree canopy for perching and territorial
activities, then a wide space, more height from the ground level and bright light for
successful mating. Although a number of mating behavioral acts in S. epius are
similar to those of some phytophagous lycaenids, the importance of vital conditions
i.e. tree canopy, canopy height from the ground level, space and daylight conditions
essential for mating in S. epius is not reported in other species. In the field, S. epius
mate location is mainly dependent on the availability of resources i.e. density of
hemipteran infestation, occurrence of tree canopy, and bright daylight. This vital
information can be utilized in mass rearing of S. epius in captivity as well as its
conservation in the field to exploit the predator as a major biocontrol agent for
mealybugs and other hemipteran pests. Furthermore, this investigation may be useful
to induce mating in captivity for rearing of any carnivorous and phytophagous
Acknowledgements The second author acknowledges the financial assistance [F. No.33-344/2007 (SR)]
from the University Grants Commission (UGC), New Delhi, India to carry out the above work. The first
author is grateful to the Council of Scientific and Industrial Research (CSIR), New Delhi, for the award of
Senior Research Fellowship.
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... We used specimens belonging to two scientific collections, which were captured by different collectors in several localities covering many parts of Mexico, along a period of several decades. Secondly, since in many Lycaenidae species males employ a sit-and-wait mate location strategy, perching in consecutive days in the same places, at the same time of the day and returning to the same perches after inspecting flying objects (Johnson & Borgo, 1976;Alcock, 1983;Cordero & Soberon, 1990;Fischer & Fiedler, 2001;Takeuchi & Imafuku, 2005a;Takeuchi & Imafuku, 2005b;Salazar, 2011;Dinesh & Venkatesha, 2013), we also explored the hypothesis that males, due to their stationary, conspicuous and predictable behaviour, suffer higher rates of predator attacks than females. ...
... Here, using larger and more heterogeneous samples, we confirm that the probability of finding symmetrical damage in the FH area is larger in species and genera with more FH components (Tables 1 and 2, Fig. 3), providing further evidence for the role of predator deflection in the evolution of butterfly false heads and for the idea that FHs with more components are better at deceiving predators. In contrast, our hypothesis that the sit-and-wait mate location strategy employed by males of many Lycaenidae species (Johnson & Borgo, 1976;Alcock, 1983;Cordero & Soberon, 1990;Fischer & Fiedler, 2001;Takeuchi & Imafuku, 2005a;Takeuchi & Imafuku, 2005b;Salazar, 2011;Dinesh & Venkatesha, 2013) results in males being more stationary and behaviourally conspicuous than females, and thus more prone to be attacked by visually oriented predators, was not supported by the data. Males and females did not differ in the proportion of specimens with symmetrical damage (Table 2). ...
Full-text available
In many butterfly species, the posterior end of the hindwings of individuals perching with their wings closed resembles a butterfly head. This "false head" pattern is considered an adaptation to deflect predator attacks to less vulnerable parts of the body. The presence of symmetrical damage in left and right wings is considered evidence of failed predator attacks to perching butterflies. In this research, we tested the prediction derived from the deflection hypothesis that the degree of resemblance of the false head area (FH) to a real head, as measured by the number of FH "components" (eyespots, "false antennae", modified outline of the FH area and lines converging on the FH area) present in the hindwings, is positively correlated to the frequency of symmetrical damage in the FH area. We studied specimens from two scientific collections of butterflies of the subfamily Theclinae (Lycaenidae) belonging to the Universidad Nacional Autónoma de México (Colección Nacional de Insectos [CNIN] and Museo de Zoología, Facultad de Ciencias [MZFC]). We scored the presence of symmetrical damage in a sample of 20,709 specimens (CNIN: 3,722; MZFC: 16,987) from 126 species (CNIN: 78 species; MZFC: 117 species; 71 species shared by both collections) whose hindwings vary in the number of FH components, and found that, as predicted, the proportion of specimens with symmetrical damage increases as the number of FH components increases. We also tested the hypothesis that behavioural differences between the sexes makes males more prone to receive predator attacks and, thus, we predicted a higher frequency of symmetrical damage in the FH of males than in that of females. We found that the frequency of symmetrical damage was not significantly different between males and females, suggesting that behavioural differences between the sexes produce no differences in the risk of being attacked. Overall, our results provide support to the idea that the FH of butterflies is an adaptation that deflects predator attacks to less vulnerable parts of the body in both sexes.
... Of course, males may be able to discriminate the sex of flying conspecifics after they catch up with them. In some territorial butterflies, males show different flight tracks to males and females: in malemale interactions, they fly around each other, whereas, in male-female interactions, males perform a specific pursuit (Rutowski, Alcock & Carey, 1989;Rutowski, 1992;Dinesh & Venkatesha, 2013). This does not mean that males can discriminate the sex of flying conspecifics because male behaviour depends on the movement of their opponent, irrespective of sexual recognition. ...
... The pair never copulates in the air. If the female alights nearby, the male also alights there, walks to her and bends his abdomen to copulate with her (Tinbergen et al., 1972;Wickman & Wiklund, 1983;Takeuchi & Imafuku, 2005;Pinzari, 2009;Dinesh & Venkatesha, 2013;Takeuchi, 2010). A male cannot copulate with a female if she does not alight (but see Carvalho et al., 2016). ...
Full-text available
Males of various flying insects perform conspicuous aerial interactions around their mating stations. The broadly accepted interpretation of their aerial interaction is a war of attrition, where two contestants perform costly displays, and the one that reaches its cost threshold earlier gives up. The implicit but important requirement in this model is that some forces that match the intensity of display of the two contestants are necessary, and failure to enforce matching allows foul contestants that delay or stop their display to avoid paying contest costs. In addition, wars of attrition require flying insects to distinguish the sex of flying conspecifics because their aerial interactions begin when intruders fly into the territory. We investigated past research on the behaviour of odonates and butterflies aiming to clarify whether the two prerequisites of wars of attrition are fulfilled: (1) contestants can inflict substantial costs on nondisplaying opponents and (2) contestants can discriminate the sex of flying conspecifics. In odonates, we found an abundance of evidence suggesting that contests involve physical attack and that the ability of sexual discrimination is sufficient. Therefore, wars of attrition may occur in territorial odonates. In butterflies, however, we could not find any evidence that the two prerequisites are filled. The aerial interactions of butterflies are better interpreted as courtship between sexually active males (the erroneous courtship hypothesis). Based on these results, we discuss future directions of research on the aerial contests of flying insects.
... Studies on an Indian Apefly species have indicated that the butterfly can be used for bio-control against Mealy bugs [18,68,86,20] . The newly hatched larvae of S. epius have been reported as capable to consume eggs while inside the ovisac of the Mealy bug [82] . ...
Full-text available
This review compiled published information on a rare butterfly, namely Apefly (Spalgis spp) in order to conserve it and explore its potential for managing papaya mealybugs (PM) (Pracoccus marginatus Williams and Granara de Willink), which attack various crop varieties of economic importance worldwide, causing a yield loss of up to 100% in Sub Saharan Africa. One of the biggest challenges behind PM management is its ability to form a wax coat that makes it difficult for chemicals used in pest management to suppress the PM once they have attacked the crops. This review provides a comprehensive description of different Spalgis species, their distribution and how they can be used to control PMs to reduce the losses of crops in Sub Saharan Africa. About 29% of the cited literature in this review indicates the Indian Apefly (Spalgis epius) to be intensively studied with explicit information of its biological control potential. The African Apefly (Spalgis lemolea) has been mentioned by 16.8% of the cited literature with limited or without detailed information ot its biological control potential. Thus, this review recommends research on understanding the biological control potential and other key information such as the life cycles, biology, diversity and ecology of the African Apefly so that the information that will be obtained can be used to design strategies towards conservation of Apefly (Spalgis lemolea. Druce) and biological management of papaya mealybug in Sub Saharan Africa. Keywords: Papaya mealybugs, Bio-control, Butterfly, Pest, Host range, Species conservation
... An ethogram is the description of an animal's behavior repertoire that forms the basis of ethological studies. Ethograms have been constructed for many animals, including insects (Dinesh and Venkatesha 2013), particularly social insects such as bees (Seeley 1982), ants (Jayasuriya and Traniello 1985), and termites (Rosengaus and Traniello 1991). In social insects, ethograms are particularly relevant in understanding division of labor among individuals in a colony, where caste polyethism and age polyethism can result in optimized growth and fitness for the colony (Seeley 1982). ...
Full-text available
A description of termite biology, distribution and diversity, economic importance, and sustainable management is presented. Liquid termiticide injection to soil, to establish a toxic or repellent chemical barrier against termites, is a traditional method applied for control. Baiting programs have been used successfully to eliminate subterranean termite colonies. Biological approaches along with entomophagy are also effective to manage termite population.
... In this case, males usually try to impress the females to mate in order to continue their generation. According to the pre-mating behavior butterflies show, they are of two types: perching species and patrolling species (Kohatsu et al. 2011;Dinesh and Venkatesha 2013 ...
Butterflies are a biodiversity indicator group and as such their presence in a certain area can be an indicator of the quality of the habitat. Like other behaviors, mating behavior of butterfly is very specific and depends on different types of aspects. In the study period of 12 months, pre-mating of 88 species (429 times) and mating of 26 species (65 times) of butterflies had spotted. Lycaenidae (7 species) was the most dominant and Nymphalidae (1 species) was the least dominant families in case of mating occurrences, other observed families were: Hesperiidae (6 species), Satyridae (4 species), Papilionidae (3 species), Pieridae (3 species) and Danaidae (2 species). Pre-mating success of all these families of butterflies was very low. Different types of pre-mating and mating behavior including perching behavior, patrolling behavior, territoriality, hair pencil behavior and pupal mating spotted during this period. Interestingly, in case of both pre-mating and mating, butterflies mostly preferred March, August and November and the relation is strongly statistical significant among each other. Butterflies are very sensitive biota and a little change in the weather can cause huge change. In this study, pre-mating and mating occurrences of butterflies were also strongly statistically significant with humidity and precipitation. The very high significant level indicates that probably, environmental condition was very critical during this study period and more research is needed to conclude. During pre-mating and mating, butterflies showed their preference towards different vegetation layers and also towards specific time schedules of a day.
... An ethogram is the description of an animal's behavior repertoire that forms the basis of ethological studies. Ethograms have been constructed for many animals, including common marmosets (Stevenson and Poole 1976), rhesus monkeys (Sade 1973), horses (Arnold and Grassia 1982), and insects (Kasuya 1983;Dinesh and Venkatesha 2013), particularly social insects such as bees (Seeley 1982;Kolmes 1985), ants (Wilson and Fagen 1974;Wilson 1976;Herbers and Cunningham 1983;Jayasuriya and Traniello 1985) and termites (Rosengaus and Traniello 1991). In social insects, ethograms are particularly relevant in understanding division of labor among individuals in a colony, where caste polyethism and age polyethism can result in optimized growth and fitness for the colony (Oster and Wilson 1978;Seeley 1982). ...
Juvenile colonies of Coptotermes formosanus Shiraki were investigated to determine the social interactions among all individuals near the central nest of a colony. The behavioral repertoire of whole colonies of subterranean termites has yet to be identified because of their cryptic nests. Colonies were placed in planar arenas, and their behavioral repertoire and activities were recorded with video cameras. All castes and larval instars were determined and behavioral interactions were monitored and described, including feeding behaviors, grooming, nest maintenance, and inactivity. When two termites interacted, it was also determined which one was the donor or the recipient of the act. An ethogram was constructed by calculating relative frequency of each behavior for first instar larvae, second instar larvae, first instar workers, second instar workers or older, queen, king, soldiers and presoldiers. Larval instars, primary reproductives and soldiers were mostly passive during interactions. Workers performed most of the tasks; however, there was a difference of task divisions among worker instars. Young workers (W 1) performed most of the grooming of individuals in the central nest, while older workers (W 2+) provided nest maintenance and sanitation, by collecting fecal matter from most nest mates. In addition, older workers were mostly in charge of caring for the primary reproductives and the maintenance of the royal chamber. This study identified instar-specific activities and provides a first insight into caste and age polyethism in C. formosanus.
Spalgis epius is an economically important hemipterophagous butterfly. Detailed information on the population dynamics, natural enemies, and prey range of S. epius and its association with mealybug-attendant ant species is lacking. Three years of field studies conducted at Bangalore University campus, Bengaluru, India on these aspects indicated that the population density of S. epius was greatest from June to December and least from February to May in the low land region. S. epius survived on eight prey species, which were present in different months, of which Phenacoccus indicus was recorded as a prey of S. epius for the first time. Different prey species occurred on 12 species of host plants. The occurrence of S. epius was negatively correlated with temperature and positively correlated with relative humidity and prey populations. Six mealybug-attendant ant species were associated with the larvae of S. epius. The seven general predators of S. epius adults were recorded. Knowledge on the population dynamics and a prey range of S. epius and its interaction with mealybug-attendant ant species could be helpful to using this predator as a major biocontrol agent of various species of mealybugs. This study contributed to our understanding of the population dynamics of a hemipterophagous butterfly.
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Many species of mealybugs (Hemiptera: Pseudococcidae) are serious pests of economically important crops worldwide. We evaluated the influence of constant temperatures: 14, 16, 18, 20, 22, 24, 26, 28, 30, 32 and 34°C on the life history and demographic parameters of Spalgis epius (Lepidoptera: Lycaenidae), a candidate biological control agent of various species of mealybugs. No eggs completed their development at 14 and 34°C. Egg-to-adult developmental time significantly decreased from 89.9 days at 16°C to 20.4 days at 32°C. The estimated lower temperature threshold of 10.2°C and 416.6 degree-days were required to complete egg-to-adult development. The mortality of immature stages was maximum at 16 and 32°C and minimum at 28°C. The highest lifetime fecundity was recorded at 28°C and it significantly decreased at 32°C. The longevity of adults was about three times more at 16°C than at 30 and 32°C. The net reproductive rate (R 0) significantly increased with increased temperatures up to 28°C and significantly decreased at 32°C. The mean generation time (T) significantly decreased with increased temperature up to 30°C, but it significantly increased at 32°C. The intrinsic rate of population increase (R m ) was highest at 30°C. The finite rate of increase (λ) was significantly greater at 30°C than at other temperatures. These data suggest that S. epius can develop, reproduce and survive in a wide range of temperatures and thus could be regarded a potential biological control agent of mealybugs.
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In this study, we examined the oviposition behavior and preference of Spalgis epius, a potential predator of mealybug crop pests. An ethogram of oviposition behavior was constructed based on observations made in an oviposition cage. Ovipositional behavioral acts were catalogued and separated into two behavioral repertoires: searching and egg laying. Gravid females of S. epius oviposited similar numbers of eggs on three mealybug species. Females preferred eggs and adults to nymphs of mealybugs for oviposition. Among three species of mealybugs attended by ants, females laid fewer eggs in the mealybug mass attended by Oecophylla smaragdina than on mealybugs attended by Tapinoma melanocephalum and Camponotus variegatus. Females preferred mealybug masses already containing conspecific eggs to mealybug masses containing conspecific larvae or Cryptolaemus montrouzieri larvae for egg deposition. Gravid females laid larger numbers of eggs under bright sunlight than in diffused sunlight or shade. The results of this study showed that S. epius can effectively attack any species of mealybugs, avoid intra- and interspecific competition, and co-exist with some species of ants attending mealybugs. With the knowledge of these behaviors, this predator can be effectively utilized as a major biological control agent of mealybugs.
The male of Papilio xuthus, a Japanese black-and-yellow Swallowtail butterfly visually determines his conspecific mate during courtship. Field experiments with various dummies and models revealed that the visual releaser of the male's approach is not the color of the wing alone but its striped pattern of black and yellow. © 1975, JAPANESE SOCIETY OF APPLIED ENTOMOLOGY AND ZOOLOGY. All rights reserved.
Male black swallowtails, Papilio polyxenes, regulate thoracic temperatures in the field by behaviorally changing abdomen position, wing position, orientation to the sun, perching duration and perching height. At relatively low ambient temperatures (22@?), postures with abdomens lowered in the shade of the wings were associated with shorter perch durations, greater flight frequencies, and higher perch heights. Between 14@? and 22@? ambient, thoracic temperatures were regulated between 28@? and 32@?. Shivering is not capable of elevating temperatures above ambient as much as basking alone does, and is seen only in disturbed individuals under conditions too cool for flight. Overheating due to metabolic heat production in flight is avoided by gliding or soaring with reduced wingbeat frequency. Swallowtails survive air temperatures above 50@? for 30-min intervals in water-saturated air, but die above 46@? at lower humidities. Swallowtails survive cooling to below 0@?. Thoracic temperatures above 24@? are apparently necessary for flight, and vigorous flight occurs only when thoracic temperatures exceed 28@?. Thoracic-abdominal temperature differences were greatest at low ambient temperatures where thoracic temperatures necessary for flight are difficult to attain or maintain. At low ambient temperatures, decreased transfers of heat from thorax to abdomen via hemolymph circulation are associated with raised abdominal postures. At higher ambient temperatures, increased circulatory exchange between abdomen and thorax (augmented by abdominal contractions and a lowered abdomen) serves to dissipate heat from the thorax, especially when the abdomen is shaded by the wings.
Behavioral thermoregulation is well studied in reptiles but has not previously been known to exist in butterflies, where it seems to be widespread and perhaps universal. Like reptiles, butterflies are heliotherms, deriving their heat almost exclusively from the sun. For reception of heat they make much use of their wings. Blood circulates in the wings, making them effective heat exchangers whose efficiency is improved by modifications of structure, color and pattern. Behavioral acts (often group-specific) for heat gain include dorsal, lateral and body basking, and ground contact. Behavioral devices for heat loss include shade seeking and probably normal and excessive respiration (evaporative cooling). In optimal air temperatures a series of minor adjustment devices permits normal activity with little concession to temperature control: sum minimizing, irradiation balancing, and wing opening. Five major thermoregulatory behavior patterns can be distinguished: minimal activity, matutinal warming, vesper warming, vernier control and cooling dominant. One or another of these is in operation at all times. Butterflies and reptiles share many behavior devices but moths differ considerably from both. They are myothermus, their major heat source being muscular energy. Heliothermic butterflies and myothermic moths are compared and contrasted with respect to the ecological significance of these specializations, particularly in feeding habits. Some of the many remaining problems are discussed, most important of which is the necessity for obtaining actual body temperatures to amplify the observational data of the present paper.