Department of Agricultural Entomology, University of Agricultural Sciences, GKVK, Bangalore , India. 2

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1 Entomologia Generalis, Vol. 35 (2014), Issue 2, Stuttgart, July 2014 Article Behavioural studies on Shoot and Fruit Borer, Conogethes punctiferalis, Guenée, (Crambidae: Lepidoptera) host-associated populations reveal occurrence of cryptic species Shashank R. Pathour 1 *, Akshay Kumar Chakravarthy 1, R. Chandrashekharaiah 1 & K.R.M. Bhanu 2 1 Department of Agricultural Entomology, University of Agricultural Sciences, GKVK, Bangalore , India. 2 Bio-Control Research Laboratories, A Division of Pest Control (India) Pvt. Ltd., 36/2, Sriramanahalli. Nr. Rajankunte, Dodballapur road, Bangalore , India. With 7 figures and 1 table Abstract: Conogethes punctiferalis, Guenée, reared on castor (Ricinus) and cardamom (Elettaria) was observed for moth emergence pattern, calling and mating behaviour and effect of interbreeding on offsprings. Females C. punctiferalis reared on castor emerged four hour (17.78 %) after lights off (ALO), those reared on cardamom emerged an hour (23.46 %) ALO. The calling frequency was more pronounced in female C. punctiferalis reared on castor compared to that on cardamom. C. punctiferalis moths reared on cardamom showed peak mating between 4 to 6 hours ALO, while 6 to 9 hours in females reared on castor. Failure of hybridization between C. punctiferalis reared on castor and cardamom suggest that the two C. punctiferalis populations segregated into two species. C. punctiferalis is a complex of species and observations, experiments, history and ecology support separation of C. punctiferalis on castor and cardamom into two. Keywords: Conogethes punctiferalis, Guenée, 1854 Castor Cardamom calling frequency hybridization cryptic species *Corresponding author: spathour@gmail.com 2014 E. Schweizerbart sche Verlagsbuchhandlung, Stuttgart, Germany DOI: / /2014/ /14/0047 $ 3.25 egt_35_2_0027_0039_pathour_0047.indd 27 6/25/ :11:18 AM

2 Shashank R. Pathour et al. 1. Introduction The diversity of herbivorous insects on earth is supposed to be a consequence of specialized plant feeding (Bernays & Chapman 1994). These phytophagous insects are often monophagous or oligophagous or polyphagous in relation to distribution of their hosts (Strong et al. 1984, Mitter et al. 1988, Farrel 1998). These adaptations to the host plants have led to differentiation in life-history traits between populations using divergent host plant species. Thus, host race formation is an intermediate step in ecological speciation without geographic isolation (Bush 1969, Dres & Mallet 2002, Howard & Patrik 2005). The behavioural adaptations of such monophagous, oligophagous or polyphagous insects are highly specialized to natural host plant species. Therefore, the question of origin of host races is addressed, with the initial step of documenting the behavioural changes considered as premating barriers. In this study, an effort to understand the differences in premating and mating mechanisms in shoot-and-fruit borer Conogethes punctiferalis suspected to be different host races have been explored (Chakravarthy et al. 1991). C. punctiferalis, commonly called as yellow peach moth, cardamom stem borer or castor shoot and fruit borer is distributed in tropical Asia, East Asia and Australia. Larvae of this moth are considered polyphagous attacking more than 120 wild and cultivated plants viz., peach, chestnut, durian, citrus, papaya, ginger, eggplant and maize worldwide (Sekiguchi 1974). In India, Castor (Ricinus communis L. (Euphorbiaceae)) is a non edible oilseed grown as major dryland crop. Cardamom, Elettaria cardamomum Maton (Zingiberaceae) is a large perennial, herbaceous, rhizomatous, monocot grown as a major spice crop in hilly areas of Western Ghats (Thimmarayappa 1996). C. punctiferalis is considered as a major pest on the two crops incurring upto 63 % yield loss in castor and more than 20 % in cardamom (Kapadia 1996). According to the unpublished reports of The Food and Environment Research Agency (FERA, UK) larvae of C. punctiferalis have been detected inside tropical fruits 18 times in the last 5 years at three points (in England and Wales) between 2007 and Due to report of damage to apple in North China (Cabi 2011), interceptions of the pest in fruits from Pakistan and a pest concern to many countries like New Zealand, South Africa, Canada and the USA, a Rapid Pest Risk Analysis on Conogethes was conducted. C. punctiferalis being a highly polyphagous pest, larvae attack fruits, seeds and stem of diverse plants. It is an important quarantine pest. Till date, C. punctiferalis infesting castor and cardamom is considered as same species. But the differences in habitat of castor and cardamom and morphological characters of C. punctiferalis infesting two hosts, elicits a doubt that C. punctiferalis populations infesting castor and cardamom may be different. To accomplish the purpose of this study, phenological differences like adult emergence pattern, calling behaviour and mating behaviour were recorded. In addition, interbreeding studies between two Conogethes populations were conducted to unravel if the population is dissimilar genetically. egt_35_2_0027_0039_pathour_0047.indd 28 6/25/ :11:18 AM

3 Behavioural studies on Shoot and Fruit Borer 2. Material and Methods 2.1. Laboratory culture of Conogethes Cardamom Conogethes pupae collected from the infested cardamom suckers from Mudigere (12 25 N, E and 980 m AMSL), Karnataka, were maintained at Behaviour Testing Lab, Bio-Control Research Laboratories (BCRL) (Pest Control India Pvt. Ltd. (PCI)) for adult emergence (25 ± 2 ºC, 80 % RH and 14:10 LD). A pair of freshly emerged moths, were sex separated based on their body size (females are bigger than male and males moths with tuft of black hairs at the abdominal tip) and released into rearing cages ( cm). Cage was provided with 10 % honey as a food source and young cardamom (< 1 year old) shoots in a conical flask (500 ml) (24 ± 5 o C, 75 ± 5 % RH, 13:11 LD) for oviposition. Fresh food was offered once in two days. Fig. 1. Daily adult emergence rhythms of C. punctiferalis reared on cardamom (n = 81) (lights off at 18:00 and on at 7:00) Castor Field collected C. punctiferalis reared on castor capsules (R. communis) in plastic trays ( cm) at Department of Entomology, University of Agricultural Sciences, GKVK, Bangalore for adult emergence. Daily emerged adults were sexed and kept in large cages ( cm) for oviposition. Cotton wicks soaked with 10 % honey solution were provided as a source of energy to the moths. In all cages, castor inflorescence (panicle) with flowers and immature capsules were provided as an ovipositional substrate. For all cages, males and females, four of each were released in to the cage and observed for mating and oviposition. Newly hatched first instar larvae were transferred to immature castor capsules and reared for adult emergence. New castor capsules were given as food once in four days (Nayar et al. 1976). egt_35_2_0027_0039_pathour_0047.indd 29 6/25/ :11:18 AM

4 Shashank R. Pathour et al. Fig. 2. Daily calling activity of female Conogethes species reared on castor (n = 42) and cardamom (n 2.2. Adult emergence When sex separated pupae turned into dark color, observed for moth emergence. Date records were maintained on emerging adults. To detect daily emergence pattern, the number of moths emerged were recorded at hourly intervals for the entire day. During the scotophase, the observations were made using a torch covered with red cellophane. A Mann-Whitney two sample two tailed rank test (Zar 1999) was used to determine whether daily emergence differed between sexes. The mean emergence time for males and females C. punctiferalis reared on castor and cardamom were quantified and analysed using ANOVA. Table 1. Comparison of the mean ages of mortality of C. punctiferalis on castor and cardamom. Age of mortality days post-eclosion (Mean ± SE) t-test P between castor and cardamom Sex Castor Cardamom Female 8.11 ± ± Male 6.97 ± ± t-test P between sexes Adult longevity Freshly emerged adult moths (male and female) were released into the rearing cages and observed for longevity. For this, males (25) and females (25) Conogethes moths egt_35_2_0027_0039_pathour_0047.indd 30 6/25/ :11:18 AM

5 Behavioural studies on Shoot and Fruit Borer reared on castor (n = 50) and cardamom (n = 50) were released into the rearing cages separately and provided with 10 % sugar syrup in a cotton wick. Moth activities were observed daily and number of days moths survived were recorded. Data were analyzed using student t-test (Zar 1999) Calling Behaviour Newly emerged female C. punctiferalis from castor and cardamom were transferred separately into transparent plastic cages ( cm). Cotton wicks with 10% honey were replenished every day. Fresh castor capsules and cardamom leaves (2 leaves/cage) were also placed in to the cage to enhance calling behavior. Female moths were examined every 15 min. for initiation and termination of calling behavior. Female was considered to be calling if their abdomen was raised or curved ventrally (Konno et al. 1980, Nakono et al. 2012). The observation like number of females expressed the calling posture was recorded at hourly intervals from 1 hour after lights off (18:00 p.m.) till lights on at (7:00 a.m.). During scotophase, observations were facilitated using a dim red colour spot light torch. Observations on pattern and number of females calling were quantified and analysed using ANOVA after effecting square root transformation. Data on duration of calling between Conogethes reared on castor and cardamom were subjected to Paired T test. Polynomial trend-line was drawn to fit the daily calling activity of female Conogethes reared on castor and cardamom. Fig. 3. Comparative mean duration of calling behaviour (mean of seven days) response of Conogethes species reared on castor and cardamom under laboratory conditions (values are means ± SD). Values followed by the same letters in each component are not significantly different (Tukey-Krammer s test, p < 0.05). egt_35_2_0027_0039_pathour_0047.indd 31 6/25/ :11:18 AM

6 Shashank R. Pathour et al Mating and Inter-breeding experiments Pupae of each population were separated by sex and observed for adult emergence. Virgin male and female moths reared on castor were paired with virgin female and male moths of C. punctiferalis reared on cardamom in the cage ( cm). Insects were used in different combinations (castor female with cardamom male, castor female with castor male, cardamom female with castor male and cardamom female with cardamom male). The choice and no-choice mating experiment was also conducted using a single male with a single female at a time or single male with multiple females. Likewise all possible combinations were tested. Five pairs of their respective population were tested simultaneously as a control. The eggs deposited on the substrate (respective host) were collected daily and hatchability rate was estimated. 3. Results 3.1. Adult emergence Man-Whitney rank test at one degree of freedom indicated non-significant differences (P > 0.01) in moth emergence pattern of C. punctiferalis species reared on castor and cardamom. Peak emergence time varied significantly (F 12,36 = 14.44, P < 0.01) among male and female C. punctiferalis reared on different hosts. In male moths reared on castor emergence period for male was 3rd h after lights off. Where as in females 4 h after lights-off. The peak emergence of males and females was non-significant (P > 0.01) throughout the scotophase. C. punctiferalis reared on cardamom found peak emergence period in 3rd and 1st h after light-off in male and female, respectively. But, their peak emergence periods was overlapping with each other and were non-significant (P > 0.01). Comparative emergence patterns of male and female C. punctiferalis bred on castor and cardamom were non-significant. The peak emergence of males in both case were 3 h after lights off. Where as in female was 2nd h after light-off (43.21 and % of moths emerged from the pupae reared on cardamom and castor, respectively). The peak hour of emergence in female C. punctiferalis reared on castor was 4 h after lights off (17.78 %) which was delayed by 3 h in peak emergence of moths reared on cardamom (1 h after lights off %) Adult longevity Observations revealed that there were non-significant statistical differences in longevity between the males and females of C. punctiferalis bred on castor and cardamom (P > 0.05). However, significant differences (P < 0.001) were evident between C. punctiferalis reared on castor and cardamom. Female moths reared on cardamom had lifespan of 8.62 ± 0.27 days (Mean ± SE), where as in castor female s 8.11 ± 0.27 egt_35_2_0027_0039_pathour_0047.indd 32 6/25/ :11:18 AM

7 Behavioural studies on Shoot and Fruit Borer days. Male moths survived 6.97 ± 0.31 days and 7.59 ± 0.26 days in C. punctiferalis reared on castor and cardamom, respectively. Adult females have long lifespan than males Calling Behaviour Castor C. punctiferalis The calling positions of female were either posing the abdomen in a curved manner or maintained the abdomen straight in a horizontal position with wing buzzing occasionally. During calling position, ovipositor was extruded beyond the abdominal tip. Apart from these, the females were moving their antennae frequently up and down and with little walking. Calling behaviour was observed in one to seven days after emergence of female moths. The onset and ending of calling behaviour was not differing significantly with the age (P > 0.001). Calling behaviour, regardless of female age, was recorded from the 1st to 11th h of scotophase, with the peak calling at 6th to 8th h after light-off (F 6,66 = 3.57, P = 0.003). Per cent calling females also increased significantly with age (F 11,66 = 20.66, P < 0.001). The mean duration of calling differed significantly with age in castor female (F 6,240 = 34.7, P < 0.00). Duration of calling was appreciably low in one day old females (P > 0.05) compared to others. Length of calling increased considerably on second day. However, on 3rd, 4th and 5th day females, even though increased length of calling evident but not significant. As days advanced (6th and 7th d), length of calling decreased significantly (P > 0.05), but still, significantly higher than the 1st and 2nd day-old females. Fig. 4. Temporal distribution of mating throughout the scotophase of Conogethes species reared on castor and cardamom (n = 30 pairs). egt_35_2_0027_0039_pathour_0047.indd 33 6/25/ :11:18 AM

8 Shashank R. Pathour et al Cardamom C. punctiferalis The calling behaviour was influenced by the age of the female (F 11,66 = 23.20, P < 0.001). There was an increased in per cent calling female throughout the scotophase and decreased thereafter (F 6,66 = 9.92, P < 0.001). Peak calling was observed on 5th to 7th h after light-off. In cardamom female, the length calling (F 6,210 = 43.00, P < 0.001) was varied significantly with age. The calling duration of cardamom female was similar to that of castor female Comparison of calling duration of castor and cardamom female In both the population, mean duration of calling gradually increased up to 4 h after light off. Subsequently, the duration of calling decreased. The length of calling in cardamom female in 3 to 4 h after lights off was significantly high (P < 0.05) compared to others. But on castor, duration of calling was significantly high (P < 0.05) 2 h after lights off. Conogethes moths reared on castor had lower mean duration (min) of calling compared to cardamom (t = 1.92, df = 34, P < 0.05). The duration of calling in moths reared on castor varied from 38.5 ± 8.91 min. to ± min. compared to ± 6.61 min. to ± min. on cardamom. Aforementioned observations established differences in pattern of calling behavior between moths reared on castor and cardamom Mating behaviour The courtship behaviour began when males flew near a calling female (within 10 cm radius) and walked towards her while rapidly vibrating his wings. Occasionally, a female would fly off when approached. The beginning of copulation always occurred between 4 and 7 h, corresponding to the interval from 180 to 360 min after dusk. The mean copulation duration was 35 min. C. punctiferalis reared on castor and cardamom had single mating. The maximum percentage of moth mated reared on cardamom was between 4 to 6 h after lights off with the peak activity in h (y = x x x x , R 2 = 0.84). This was much earlier (peak mating in h) than observed in moths reared on castor (6 to 9 hours) (y = x x x x , R 2 = ). But, there was no statistical difference in their peak mating time between castor and cardamom females. The per cent mated were lower (maximum %) in Conogethes on castor compared to that on cardamom (maximum %). This observation suggests that maximally per cent of moths mated under laboratory conditions when reared on castor and cardamom Interbreeding experiments The mating experiments were conducted in two sets viz., single pair/cage and multiple pairs/cage. This is to confirm whether single or multiple pairs influence mating and egt_35_2_0027_0039_pathour_0047.indd 34 6/25/ :11:19 AM

9 Behavioural studies on Shoot and Fruit Borer mating success. Flowchart in Figs. 5 and 6 indicated results of inter and intra cross mating experiments between C. punctiferalis reared on castor and cardamom with a pair/cage. Fig. 5. Flow chart indicating results of inter and intra cross mating experiments between Conogethes reared on castor and cardamom (single pair/cage). Fig. 6. Flow chart indicating results of inter and intra cross mating experiments between Conogethes reared on castor and cardamom (multiple pair/cage). The moths were maintained under standardized condition developed for C. punctiferalis rearing. Moths reared on castor recorded 25 % mating (Fig. 7A) success with mean number of eggs / female at with 68 % hatching. Similarly, when moths egt_35_2_0027_0039_pathour_0047.indd 35 6/25/ :11:19 AM

10 Shashank R. Pathour et al. were reared on cardamom % mating (Fig. 7B) success was recorded with eggs/female and with % hatching when a pair / cage. When moths reared on two different hosts were crossed, no mating occurred although the moths attempted to mate, several times. A Fig. 7. Mating pair of Conogethes species breeding on A) castor and B) cardamom. B In case of multiple pairs / cage, % mating success with mean number of eggs/ female at with %. Similarly, when moths were reared on cardamom % mating success was recorded with eggs/female and with % hatching. Data of multiple pairs / cage were similar to single pair/cage and no successful mating occurred although the moths attempted to mate, several times. Thus, the failure of hybridization between Conogethes bred on castor and cardamom suggest that they might have segregated into two different species. egt_35_2_0027_0039_pathour_0047.indd 36 6/25/ :11:19 AM

11 Behavioural studies on Shoot and Fruit Borer 4. Discussion Moth emergence rhythm of laboratory reared Conogethes on castor and cardamom revealed that the males and females emerged during scotophase. This is the first attempt to study the emergence pattern in detail on Conogethes reared on different hosts. In moth emergence studies, female C. punctiferalis reared on cardamom showed peak emergence earlier (Peak emergence 1 h of scotophase) than those of female reared on castor (Peak emergence 4 h of scotophase). Similarly, in male moth also the peak emergence period was not overlapping, when they were reared on different hosts. In C. punctiferalis, the daily emergence rhythm of female moths was periodic and not synchronized when they were reared on different host plants. The adult longevity of moths reared on castor and cardamom males and females was significantly superior under laboratory conditions. Moths reared on cardamom live longer than those of moths reared on castor. The reasons for differences in their longevity were unknown. Similar studies were conducted by Bilapate & Talati (1978) on C. punctiferalis reared on castor and Thyagaraj (2003) in C. punctiferalis breeding on cardamom. Results of laboratory experiments showed no difference in calling positions of C. punctiferalis females reared on different hosts. This calling behaviour is characteristic for several species. But, the abdomen bending and extension of the ovipositor to expose the pheromone glands is the main characteristic of the calling posture in C. punctiferalis reared on castor and cardamom. The female calling frequency of C. punctiferalis reared on castor was more pronounced compared to C. punctiferalis reared on cardamom with moths one to two days old. In three to seven days old moths, the peak calling in female C. punctiferalis reared on cardamom was recorded two h earlier than C. punctiferalis reared on castor. Similarly, the mean duration of calling in C. punctiferalis reared on castor peaked at 4 h after lights off compared with 3 h after lights off in C. punctiferalis reared on cardamom. The calling behavior of C. punctiferalis infesting peaches was studied by Konno (1986) reveled that calling started from 5 h after light-off, reached a maximum 7.5 h after light-off and then decreased. According to Rajabaskar & Regupathy (2012) the peak calling behaviour of C. punctiferalis females on cardamom was observed 3 h prior to end of scotophase. This investigation is not in line with the present findings, may be due to different laboratory conditions and in study conducted by Rajabaskar & Regupathy (2012) larvae were collected from cardamom fields and reared on ginger in laboratory. The change in the host in laboratory influenced calling behaviour of female moths following Hopkins s host selection principle (Hopkins 1917). Mating in Conogethes reared on castor and cardamom showed that there is a significant difference in mating time. The maximum per cent of moths mated reared on cardamom was between 4 to 6 hours compared with castor 6 to 9 h. The failure of hybridization between Conogethes populations reared on castor and cardamom suggested that they might have segregated into two species. This suggestion is based on the fact that the moths reared on the two hosts visibly look alike and branded as same species until now. Morphologically adults and larvae are similar (may not be identical), co-occuring in a habitat. For instance, in cardamom egt_35_2_0027_0039_pathour_0047.indd 37 6/25/ :11:19 AM

12 Shashank R. Pathour et al. growing tracts of South and North East India, the inhabiting area of two Conogethes populations overlap. Both Conogethes populations principal host plants overlap (at least in Orient). Centre-of-origin of both the Conogethes populations lies in the Indian subcontinent. Distribution of castor extend from oriental to south east Mediterranean to East Africa, of Cardamom from Asia to Australia (Philips & Rix 1999) and of C. punctiferalis from Asia to Australia (Cabi 2011). Thus, Conogethes represent a complex of species and incomplete information on their separation has been published with Robinson (1994) suggesting that there may be around 20 species in the genus. This is in line with the species definition of Mayr (1969). Studies of Konno et al. (1981) revealed that the two types of Dichocrosis (= Conogethes) punctiferalis viz., fruit feeding type and Pinaceae feeding type have not shown any mating suggesting the sexual isolation. Studies have been conducted on other crambids by Langille & Keaste (1972), Liebherr & Roelofs (1975), Seol et al. (1986) and Konno & Tanaka (1996) with similar conclusions. Thus, observations on female moth emergence, differences in female calling behaviour, absence of hybridization and differences in adult longevity implied that the two populations were genetically different and reproductively isolated. Acknowledgements We thank Dr. C.A. Virakthamath, University of Agricultural Sciences, Bangalore, India and Dr. Hiroshi Honda, Graduate School of life and Environmental Sciences, Tsukuba, Japan for valuable contributions. This work was supported by Department of Science and Technology (DST), New Delhi by awarding the INSPIRE fellowship. References Ayar, K.K., Ananthakrishnan, T.N. & David, B.V. (1976): General and Applied Entomology. Tata McGraw Hill, New Delhi. Bernays, E.A. & Chapman, R.F. (1994): Host-Plant Selection by Phytophagous Insects. Chapman and Hall, New York, USA. Bilapate, G.G. & Talati, G.M. (1978): Some studies on bionomics of castor short and fruit borer Dichocrosis punctiferalis from Gujarat. J. Maharashtra Agri. Uni. 3 (1): Bush, G.L. (1969): Sympatric host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae). Evolution 23: Cabi (2011): Conogethes punctiferalis datasheet. Crop Protection Compendium, Wallingford, UK-CAB International. Chakravarthy, A.K., Honda, H. & Thyagaraj, N.E. (1991): Comparison of containers for larval rearing in stalk and fruit feeding type of Conogethes punctiferalis (Guen.) (Lepidoptera: Pyralidae). Placrosym 9: Dres, M. & Mallet, J. (2002): Host races in plant-feeding insects and their importance in sympatric speciation. Philos. Trans. R. Soc. Lond. B 357: egt_35_2_0027_0039_pathour_0047.indd 38 6/25/ :11:19 AM

13 Behavioural studies on Shoot and Fruit Borer Farrell, B.D. (1998): Inordinate fondness explained: why are there so many beetles? Science 281: Hopkins, A.D. (1917): A discussion of C. G. Hewitt s paper on Insect Behavior. J. Econ. Entomol. 10: Howard, D.R. & Patrik, N. (2005): Ecological speciation. Ecol. Lett. 8, Paris. Kapadia, M.N. (1996): Estimation of loses due to pod borer in oil seed crops. J. Oil Seeds Res. 13 (1): Konno, Y. & Tanaka, F. (1996): Mating time of the rice-feeding and water-oat-feeding strains of the rice stem borer, Chilo suppressalis (Walker) (Lepidoptera: Pyralidae). Japan. J. Appl. Entomol. Zool. 40: Konno, Y. (1986): Time-lag between sex pheromone content and the calling behavior in the yellow peach moth, Conogethes punctiferalis (Guenee) (Lepidoptera: Pyralidae). Appl. Entomol. Zool. 21: Konno, Y., Honda, N. & Matsumoto, Y. (1980): Observation on the mating behaviour and biology for the six pheromone of yellow peach moth Dichocrosis punctiferalis. Appl. Entomol. Zool. 15 (3): Konno, Y., Honda, N. & Matsumoto, Y. (1981): Mechanism of reproductive isolation between the fruit feeding and pin ace feeding type of yellow peach moth, Dichocrosis punctiferalis. Japan. J. Appl. Entomol. Zool. 25 (4): Langille, R.H. & Keaste, A.J. (1972): Mating Behavior of the Southwestern Corn Borer. Environ. Entomol. 2 (2): Liebherr, J. & Roelofs, W.L. (1975): Laboratory hybridization and mating period studies using two pheromone strains of Ostrinia nubilalis. Ann. Entomol. Soc. Amer. 68: Mayr, K. (1969): Principles of systematic zoology. McGraw-Hill, NewYork. Mitter, C., Farrell, B.D. & Wiegmann, B. (1988): Phylogenetic study of adaptive zones: has phytophagy promoted insect diversification? Amer. Nat. 132: Nakano, R., Takanashi, T., Ihara, F., Mishiro, K., Toyama, M. & Ishikawa, Y. (2012): Ultrasonic courtship song of the yellow peach moth, Conogethes punctiferalis (Lepidoptera: Crambidae). Appl. Entomol. Zool. 47 (2): Rajabaskar, D. & Regupathy, A. (2012): Calling behaviour and attractive response of cardamom shoot and capsule borer, Conogethes punctiferalis Guenee (Lepidoptera: Pyralidae) to the female crude extract and synthetic blend. J. Entomol. 2012: 1 6. Roger, P & Rix, M. (1999): Annuals and Biennials. Macmillan, London. Sekiguchi, K. (1974): Morphology, biology and control of the yellow peach moth, Dichocrosis punctiferalis Guenée (Lepidoptera: Pyralidae). Bull. Ibaraki Hort. Expt. Stn. Spec.Iss. 89. Seol, K.Y., Honda, H. & Matsumoto, Y. (1986): Mating behaviour and the sex pheromone of the lesser mulberry pyralid, Glyphodes pyloalis Walker (Lepidoptera: Pyralidae). Appl. Entomol. Zool. 21 (2): Strong, D.R., Lawton, J.H. & Southwood, R. (1984): Insects on Plants, Community Patterns and Mechanisms. Harvard Univ. Press, Cambridge. Thimmarayappa, M. (1996): Integrated nutrient management in cardamom (E. cardamomum Maton.). M. Sc. Agri. Thesis, UAS, Bangalore. Thyagaraj, N.E. (2003): Integrated Management of some important cardamom pests of hill region of Karnataka, South India. Ph. D. Thesis, Dr. B.R. Ambedkar University, Uttar Pradesh. Zar, J.H. (1999): Biostatistical Analysis. Prentice Hall Englewood Cliffs, New Jersey. Manuscript received: 11 November 2013 Accepted: 6 February 2014 egt_35_2_0027_0039_pathour_0047.indd 39 6/25/ :11:19 AM

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