Characteristics of parasitism of diamondback moth by Oomyzus sokolowksii (Hymenoptera: Eulophidae)

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1 Characteristics of parasitism of diamondback moth by Oomyzus sokolowksii (Hymenoptera: Eulophidae) N. S. Talekar Asian Vegetable Research and Development Center, Shanhua, Tainan 741, Taiwan, ROC Abstract Laboratory, greenhouse and field studies were conducted on Oomyzus sokolowskii Kurdjumov, a parasite of diamondback moth, Plutella xylostella (L.), to judge its suitability for introduction in the field to control the plutellid. Oomyzus sokolowskii preferred the third and fourth instar diamondback moth larvae over fresh pupae for parasitization. It is thus a larval parasite. Within the range of 1 C to 35 C, the higher the temperature the higher was the parasitism rate. High parasitism at temperatures of 3 C and 35 C indicates that this insect is suitable for introduction in the tropical lowlands. In a no-choice test where only fresh pupae of Cotesia plutellae Kurdjumov (another potentially competing larval parasite of diamondback moth) were offered, O. sokolowskii failed to parasitize the pupae. In a choice test where the fourth instar diamondback moth larvae and fresh C. plutellae pupae were offered, O. sokolowskii parasitized only diamondback moth larvae. This parasite, therefore, is not a hyperparasite of diamondback moth. When C. plutellae-oviposited diamondback moth larvae were offered at intervals for parasitism by O. sokolowskii, it parasitized only freshly oviposited host larvae. The longer the period that elapsed after C. plutellae oviposition of diamondback moth larvae, the lesser was the parasitism of these larvae by O. sokolowskii. In a field cage study, as the diamondback moth population increased, the parasitism of the pest by the eulophid increased, parasitism by C. plutellae, however, decreased. Host-plant (cabbage) age did not affect the parasitism of diamondback moth larvae by O. sokolowskii; in both seedlings and mature plants the level of parasitism of the plutellid larvae was comparable. Most organic insecticides tested were toxic to both pupae and adults of O. sokolowskii but Bacillus thuringiensis was not toxic. Introduction of O. sokolowskii in a large field cage erected over a cabbage field reduced the infestation of cabbage by diamondback moth and doubled the yield of cabbage over the control plot where no parasite was used. Key words: Oomyzus sokolowskii, diamondback moth, parasitism, temperature effect, hyperparasitism, parasite competition Introduction In tropical to subtropical Asia, major cruciferous vegetables are grown in two distinct ecological zones: cool highlands and hot lowlands. In both areas, diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is a serious pest. The climate, especially temperatures, in the highlands resembles that of temperate areas. In the lowlands, high temperatures reaching and at times exceeding 35 C are common. In Europe, where the diamondback moth originated, the pest is kept under control by a plethora of temperate-climate natural enemies, mainly parasites (Mustata, 1992). Introduction of some of these temperate-climate parasites, especially Diadegma semiclausum Hellen, and to some extent Diadromus collaris Gravenhorst, in countries in Asia, has resulted in their establishment in highlands resulting in considerable reduction in the severity of diamondback moth damage (Talekar, 1996a). In lowlands, however, none of these parasites is effective due to their sensitivity to high tempeatures. Cotesia plutellae Kurdjumov, which is tolerant to the high temperatures commonly found in the tropical lowlands (Talekar and Yang, 1991), occurs naturally in Taiwan (Wu, 1968), Philippines (Velasco, 1983), Malaysia (Lim and Ko, 1975) and Thailand (Keinmeesuke, 1992). Although this braconid is an important mortality factor in the lowlands, its effectiveness in controlling diamondback moth has not been consistent. Therefore, another parasite, Oomyzus sokolowskii Kurdjumov (Hymenoptera: Eulophidae), which has given very effective control of the plutellid pest in Cape Verde Island (Lima and van Harten, 1985; Carl, 1992), was imported from that tropical West African country via the International Institute of Biological Control for introduction in lowlands of Asia. Several laboratory, greenhouse, and field experiments have been conducted to study characteristics of its parasitism to judge the suitability of this parasite for introduction in tropical to subtropical areas of Asia. Preference of O. sokolowskii for various immature stages of diamondback moth Several reports indicate that O. sokolowskii is a pupal parasite (Bennet and Yaseen, 1972; Wakisaka et al. 1991; Chelliah and Srinivasan, 1986). In our first study we compared the parasitism of diamondback moth pupae with that of third and fourth instar larvae. Ten third and fourth instar larvae and freshly formed diamondback moth pupae were placed on a cabbage leaf in a 1.5-liter plastic container. Thirty 5-day-old O. sokolowskii adults were then introduced Biologically-based technologies 97

2 in the container for oviposition. We then removed the host larvae and maintained them until pupation. The numbers of larvae pupating into diamondback moth or parasite pupae were recorded. We maintained the plutellid pupae that were also exposed to O. sokolowskii for oviposition and recorded the number of pupae that produced the plutellid adults and the number that produced parasite adults. This experiment was conducted three times, each time with three replicates. In the second experiment, 2 first, second, third, and fourth instar larvae and 2 freshly formed pupae of diamondback moth were placed on large cabbage leaves inside each of four 1.5-liter round jars. Two hundred O. sokolowskii adults were then introduced in each jar and allowed 24 hours for oviposition. The number of pupae that developed into plutellid adults and the number that produced parasite adults were recorded. Host larvae were reared until pupation and the number of those pupating into diamondback moth or O. sokolowskii pupae was recorded. The rates of parasitism of immature stages from the above two experiments were analyzed by ANOVA. Mean percentage parasitism was compared by the test of Least Significant Difference (LSD). Oomyzus sokolowskii parasitized the third and fourth instar diamondback moth larvae (average 7% and 61%, respectively) but failed to parasitize pupae (Table 1). It is thus a larval parasite. Cotesia plutellae, another larval parasite of diamondback moth with similar ecological requirements as O. sokolowskii, parasitizes second, third, and fourth instar plutellid larvae with preference for the second instar (Talekar and Yang 1991). In most of southeast Asia where C. plutellae is established, the introduction of O. sokolowskii might result in competition between the two parasites. This led us to study the preference of O. sokolowskii adults to parasitize various instars of diamondback moth. Results (Table 2) indicate that O. sokolowskii prefers third and fourth instars over first and second instars. The preference of O. sokolowskii for third and fourth instars and that of C. plutellae for second instar could reduce competition between these two larval parasites, assuring survival of both. This could contribute to the biological control of diamondback moth in lowland areas of Asia where this plutellid pest is especially serious. However, the reduction of competition will depend on the interval between parasitism of second instar host larvae by C. plutellae and molting into third instar when chances of its being attacked by O. sokolowskii increases. The longer the interval, the better are the chances of survival of C. plutellae (see later discussion). Effect of temperature on parasitism Three-to five-day-old O. sokolowskii adults and third and fourth instar diamondback moth larvae were separately maintained at various temperatures: 1, 15, 2, 25, 3, and 35 C for 6 hours to condition both host and parasite to these temperatures. Fifty Table 1. Preference of O. sokolowskii for parasitism of various immature stages of diamondback moth Test Parasitism (%) 3rd instar 4th instar Pupae larvae larvae First 66.6± ±19.5 Second 68.9± ±27. Third 75.2± ± 5.1 Mean 7.2± ±14.8 Ten larvae or fresh pupae of diamondback moth were exposed to 3 O. sokolowskii adults for oviposition. Table 2. Preference of O. sokolowskii for parasitism of various instars and pupae of diamondback moth Instar Parasitism (%) (a) First Second Third Fourth Pupae LSD (b) 2. 8 (a) Parasitism data are means + standard deviation of 4 replicates (b) LSD = Least significant difference O. sokolowskii adults were then introduced in a 15- cm-diameter, 3-cm-long acrylic cylinder containing 5 plutellid larvae feeding on a cabbage leaf. Both ends of the cylinder were covered with a single layer of fine muslin cloth. Four such cylinders containing parasite adults and host larvae were maintained at 1, 15, 2, 25, 3 or 35 C for 24 hours. During this period, O. sokolowskii adults laid eggs in diamondback moth larvae. After 24 hours, the pest larvae were maintained at 25 ± 2 C and reared until pupation. At pupation the number of larvae developing into diamondback moth pupae and parasite pupae were recorded and percent parasitism was calculated. A simple linear regression correlation between temperature and percentage parasitism was calculated (Little and Hills, 1975). As the temperature increased from 1 C to 35 C, parasitism of diamondback moth by O. sokolowskii increased significantly (r =.987, p =.1, Figure 1). High parasitism at temperatures of 3 C and 35 C indicates that this parasite might be suitable for the tropical lowlands. Parasite mortality ranged from 4.5% to 14.6% and was not related to temperature. Oomyzus sokolowskii is now being introduced in crucifergrowing lowland areas of Taiwan, Thailand, and Malaysia. Hyperparasitism study Because environmental conditions, especially temperature, for survival and multiplication of C. plutellae and O. sokolowskii are similar, and because O. sokolowskii is a minute insect, it has often been postulated but never documented that O. sokolowskii could be a hyperparasite of diamondback moth. Both no-choice and choice tests were used to study the preference of O. sokolowskii for oviposition in 98 Proceedings: The Management of Diamondback Moth and Other Crucifer Pests

3 diamondback moth larvae or C. plutellae pupae. In the no-choice test, 5 fresh C. plutellae pupae were placed on a cabbage leaf in each of eight acrylic containers. Twenty two-day-old O. sokolowskii adults were then introduced for oviposition in each of four containers. The remaining four containers were maintained as control. The host and parasite insects were maintained together at 26 ± 2 C for 48 hours. Oomyzus sokolowskii adults were then removed and C. plutellae pupae transferred individually into glass vials to observe pupation and emergence of C. plutellae or O. sokolowskii adults. In the choice test, 5 fourth instar diamondback moth larvae and 5 fresh C. plutellae pupae maintained on a cabbage leaf were placed in each of four acrylic containers. Twenty two-day-old O. sokolowskii adults were then introduced inside each of these containers and allowed to lay eggs for 48 hours. The plutellid larvae and C. plutellae pupae were then maintained separately at 25 ± 2 C. The number of diamondback moth larvae producing parasite pupae and the number developing into diamondback moth pupae were recorded. Cotesia plutellae pupae were observed and the number of those that showed parasitism by O. sokolowskii was recorded. In both choice and no-choice tests, O. sokolowskii failed to parasitize C. plutellae pupae (Table 3). In the choice test O. sokolowskii parasitized the fourth instar diamondback moth larvae but not C. plutellae pupae. It was not possible to expose larvae of C. plutellae and diamondback moth for O. sokolowskii oviposition simultaneously for fair comparison because C. plutellae larvae develop inside diamondback moth larvae. Instead we used freshly formed C. plutellae pupae. The fact that in the no-choice test O. sokolowskii did not parasitize C. plutellae shows that O. sokolowskii is not a parasite of C. plutellae and thus is not a hyperparasite of diamondback moth. It is possible that O. sokolowskii parasitizes other diamondback moth parasites such as D. semiclausum or D. collaris. However, D. semiclausum is a cool climate parasite (Talekar and Yang, 1991) and D. collaris, which also prefers cool climate, is a pupal parasite. These factors rule out O. sokolowskii being a hyperparasite of diamondback moth. This finding allows the introduction of this parasite in all lowland areas of southeast Asia where C. plutellae also parasitizes diamondback moth. Competition for parasitism between C. plutellae and O. sokolowskii Since C. plutellae and O. sokolowskii share similar ecological niches, and they both attack diamondback moth larvae, it is possible that these insects would compete for host larvae and that the less competitive of the two species might not survive. Therefore, we performed an experiment to determine competition between the two parasites. Because C. plutellae can infest all four instars of diamondback moth (Velasco, 1983; Talekar and Yang, 1991) and because it is already established in most countries of Asia, we exposed the host larvae first to C. plutellae and then to O. sokolowskii. Twenty fourth instar diamondback moth larvae were individually confined with a gravid C. plutellae female in a test tube. After oviposition, each plutellid larva was observed under the microscope to make sure that C. plutellae had indeed laid eggs in each of them. The parasite eggs could be seen through larval cuticle. At, 24, 48, 72, ans 96 hours after oviposition, the C. plutellae-oviposited larvae were individually exposed to oviposition by O. sokolowskii. We recorded the number of diamondback moth larvae pupating into C. plutellae or O. sokolowskii pupae. We also recorded the mortality of diamondback moth larvae and parasite pupae. When C. plutellae-oviposited diamondback moth larvae were immediately exposed to parasitization by O. sokolowskii, most of the diamondback moth larvae were oviposited by the latter parasite and yielded O. sokolowskii pupae (Figure 2). Even 24 hours after oviposition by C. plutellae, practically 5% of the plutellid larvae were successfully parasitized by O. sokolowskii. However, after 48 hours, practically all C. plutellae-oviposited diamondback moth became C. plutellae adults. It is possible that at 48 hours and later, C. plutellae embryos, which are in an advanced stage of development, kill O. sokolowskii eggs as they do D. semiclausum eggs (Yang et al., 1994). This implies that O. sokolowskii could become the dominant parasite when introduced in an area where C. plutellae is already established. The latest information from Cape Verde Island indicates this to be the case (Carl, 1992). However, the fact that C. plutellae can parasitize earlier instars of diamondback moth larvae should enable the parasites to co-exist, as has been shown in Table 3. Parasitism of P. xylostella or C. plutellae by O. sokolowskii in choice and no-choice test Test condition Parasitism (%) Adults emerged (%) Mortality (%) P. xylostella C. plutellae P. xylostella C. plutellae P. xylostella C. plutellae No choice (a) 98.5 ± ± 1.91 Choice (b) 3.9 ± ± ± ± ± 1. Data are means ± standard deviation of 4 replicates. (a) Only C. plutellae pupae were offered for parasitism by O. sokolowskii (b) Both 4th instar P. xylostella larvae and C. plutellae pupae were offered for parasitism by O. sokolowskii. Biologically-based technologies 99

4 Parasitism (%) Paratisim (%) by 3 2 y = x r 2 = Temperature ( C) Figure 1. Parasitism of diamondback moth larvae by O. sokolowskii at various temperatures No. larvae/plant Figure 3. Relationship of population of diamondback moth larvae per plant and parasitism by O. sokolowskii Paratisim (%) by 1 Survival rate (%)( ) 1 No. adults emerged/pupa ( ) C. plutellae O. sokolowskii Hours after oviposition by C. plutellae Figure 2. Effect of time elapsed after C. plutellae oviposition in diamondback moth larvae on the success of parasitism of the same larvae by O. sokolowskii. (Each point is the average of 2 larvae exposed to parasitism). Sticker Bt Profenofos Abamectin Cartap Cyromaszine Mevinphos Bifenthrin Figure 4. Effect of various insecticides on the survival of pupae and emergence of O. sokolowskii adults Survival rate (%) Sticker Bt Abamectin Cyromazine 4 2 Bifenthrin Profenofos Mevinphos Cartap Days after treatment Figure 5. Effect of various insecticides on the survival of adults of O. sokolowskii 1 Proceedings: The Management of Diamondback Moth and Other Crucifer Pests

5 southern Japan, India, and the West Indies (Hirashima et al., 1989, Chelliah and Srinivasan, 1986, Yaseen, 1978). Relationship between diamondback moth larval population density and parasitism by O. sokolowskii In this field experiment we studied the dependence of rate of O. sokolowskii parasitism on the density of diamondback moth larval population. In a.1-ha parcel of land planted to common cabbage we introduced 5 diamondback moth adults to initiate pest infestation. Two days later we introduced 1 O. sokolowskii adults in the same area. In addition, 5 parasite adults each were also introduced two, three, and four weeks later. Once a week we monitored the larval population of diamondback moth on cabbage plants and parasitism of the pest larvae by O. sokolowskii or C. plutellae which is already established in lowland areas of Taiwan. Diamondback moth population increased steadily from 1 days after the plutellid adult release up to harvest, from 2 to 22 per 1 plants. Parasitism of larvae by O. sokolowskii increased steadily from 1.3% to 21.9%. There was significant correlation (r =.844) between larval population density and the rate of parasitism (Figure 3). Parasitism by C. plutellae decreased from 48% to 3.1% during this period. In our earlier study we found similar decrease in parasitism of C. plutellae from the beginning towards the end of the season (AVRDC, 1992). Based on our present findings, it appears that introduction of O. sokolowskii will not compete but supplement the control of diamondback moth achieved by C. plutellae. Effect of host-plant age on the parasitism of diamondback moth by O. sokolowskii A field experiment was conducted to study the influence of host-plant (cabbage) age on the parasitism of diamondback moth larvae by O. sokolowskii. A parcel of land was rototilled and worked into 24, 1.5- m wide and 5.-m long single bed plots. The whole area on four sides was confined by 2-m high fine-mesh net. The same mesh net was used to cover the top. Once every week for eight consecutive weeks we transplanted four-week-old cabbage seedlings in three randomly selected plots; one plot per replicate. Plants were maintained free of plutellid infestation. When the cabbage plants in the last transplanted plot were in the field for one week, we introduced 1 third-instar diamondback moth larvae per plant on each of 1 plants selected at random in each plot. The plants that received the pest larvae were marked. Immediately after the introduction of diamondback moth we released 5 O. sokolowskii adults inside the cage for parasitism. At three and five days after parasite introduction, we collected 3 larvae from each plot and reared them in the laboratory until pupation. We recorded the number of O. sokolowskii and diamondback moth pupae that developed and computed the percent parasitism. Parasitism varied between 18% and 54% in the first observation (three days) and 3 and 5% in the second observation (five days). There was no statistically significant relationship between plant age and parasitism. Oomyzus sokolowskii parasitism of diamondback moth is thus not affected by host-plant age. This parasite, therefore, can be introduced in the field for the biological control of the plutellid at any stage of plant growth. Effect of various insecticides on the survival of O. sokolowskii Our experience indicates that despite best efforts by researchers and extension authorities, some farmers continue to use insecticides in crucifer growing areas where parasites are newly established and diamondback moth is no longer a serious pest. Some of the insecticide use is for controlling insect pests other than diamondback moth. It is possible that some of the insecticides used could adversely affect diamondback moth parasites and thereby exacerbate diamondback moth problem. We conducted, therefore, a laboratory experiment to study toxicity of some commonly used insecticides to O. sokolowskii. Commercial formulations of each of seven insecticides and one commonly used sticker were diluted as directed on the insecticide container for use in the field. Freshly formed pupae of O. sokolowskii were then dipped in insecticide solution for five seconds. Treated pupae were air-dried under a gentle flow of air. The dry pupae were placed at 28 ± 2 C and emergence into O. sokolowskii adults was monitored. The number of pupae that failed to emerge were considered as dead. The total number of adults that emerged from surviving pupae were recorded. In a simultaneous test with adults, the insecticide solutions were sprayed on fresh cabbage leaves. The treated leaves were placed in a 15-cm diameter acrylic cylinder. Twenty O. sokolowskii adults were released inside each cylinder and insect mortality was recorded at 72 hours after treatment. The survival rate and the number of O. sokolowskii adults which emerged from the surviving pupae are shown in Figure 4. Bifenthrin was the most toxic and Bacillus thuringiensis were the least toxic insecticides. The order of toxicity was bifenthrin > mevinphos > cyromazine > cartap > abamectin > profenofos > B. thuringiensis. For adults, mevinphos and cartap were equally toxic and B. thuringiensis was the least toxic (Figure 5). The order of toxicity was mevinphos = cartap > profenofos > bifenthrin > abamectin = cyromazine > B. thuringiensis. The sticker was nontoxic to adults and its toxicity to pupae was lower than any products included in the test. Bacillus thuringiensis was the least toxic to both adults and pupae. This biological insecticide is frequently used to control diamondback moth. Abamectin, at present the most popular chemical for the control of diamondback moth, was relatively less toxic to both stages of O. sokolowskii. Cyromazine is frequently used for the control of leaf miner. This chemical is relatively safer to O. sokolowskii adults but not to Biologically-based technologies 11

6 pupae. Mevinphos, an old broadspectum organophosphorus insecticide, is at times still used for controlling diamondback moth. This chemical is highly toxic to the parasite, as is bifenthrin, a synthetic pyrethroid. Control of diamondback moth by O. sokolowskii Prior to the introduction of O. sokolowskii for the control of diamondback moth in farmers fields, we conducted a study at AVRDC to investigate the potential of this parasite under field conditions. A parcel of land planted to common cabbage was divided into three 27 m by 13.5 m plots. Each area was subsequently confined on all four sides and the top by fine mesh nylon net to prevent movement of insects between the plots. In two plots we introduced 1 diamondback moth adults each. The third plot was maintained insect free. One week later, in one of the two plots where the pest was introduced, we released all O. sokolowskii adults emerging from 1 parasitized diamondback moth cocoons. Additional 5 parasite cocoons were each released at two, three, and four weeks later. We monitored the extent of diamondback moth parasitism throughout the season and determined yield and marketable cabbage heads at harvest. In the parasite-released cage, parasitism of the plutellid larvae by O. sokolowskii ranged from 1.28% to 21.74%. Parasitism, which was very low, hovering around 2% early in the season, increased to 21.74% two weeks before harvest. Diamondback moth larval population, which remained low (around 5 larvae/ plant) through most of the season increased to 22.4 larvae per plant a week before harvest. We are unable to determine the reason for this sudden increase in population. The rate of parasitism was obviously not adequate to prevent pest population build up. In the diamondback moth-only plot, the larval population was high and reached 36.9 larvae/plant a week before harvest. In this plot, cabbage heads, on an average, weighed 1.43 kg per head. The head weight increased to 1.53 kg in the plot where both diamondback moth and O. sokolowskii were released. In the control plot, where no diamondback moth nor parasite was released, each cabbage head weighed 1.62 kg. Only 26.1% of the heads were marketable in the diamondback mothonly plot; marketability increased to 44% in the parasite-released plot but it was below 76.1% marketability of the control plot where no insects were released. Cabbage yield was 6.84 t/ha in the only diamondback moth-plot; it increased to t/ha due to parasite introduction but it was still below the t/ha obtained in the control plot. Marketable heads, rather than head weight, contributed to increased yield. Introduction of O. sokolowskii to control diamondback moth increased head weight, marketable heads, and practically doubled the yield of cabbage. These data indicate the merits of introduction of O. sokolowskii to control diamondback moth to increase cabbage yield. References AVRDC. (1992) Progress Report. Asian Vegetable Research and Development Center, Shanhua, Taiwan, 41 pp. Bennet, F. D. and Yaseen, M. (1972). Parasite introductions for the biological control of three insect pests in the Lesser Antilles and British Honduras. PANS. 18: Carl, D. (1992). DBM control project in Togo, Benin and Cape Verde. IOBC global working group on biological control of Plutella, Newsletter 1991, 24 pp. Chelliah, S. and Srinivisan, K. (1986) Bio-ecology and management of diamondback moth in India, In Diamondback moth management: Proceedings of the first international workshop (ed Talekar N. S. and Griggs, T. D.) pp Shanhua, Taiwan: Asian Vegetable Research and Development Center. Hirashima, Y., Abe, M., Tadauchi, O., Konishi, K., and Maeto, K. (1989). The hymenopterous parasitoids of the diamondback moth, Plutella xylostella (Lepidoptera, Yponomeutidae) in Japan. Esakia 28: Keinmeesuke, P. (1992). Integrated pest management of diamondback moth on crucifers in Thailand. In Collaborative Vegetable Research in Southeast Asia: Proceedings of the AVNET-I Final Workshop and AVNET-II Joint Planning Meeting, pp Shanhua, Taiwan: Asian Vegetable Research and Development Center. Lim, G. S. and Ko, W. W. (1975). Apanteles plutellae Kurdj., a newly recorded parasite of Plutella xylostella (L.) in Malaysia. MARDI Research Bulletin 3: Lima, M. L. L. and Van Harten, A. (1985). Biological control of crop pests in Cape Verde. Current situation and future programme. Rev. Invest. Agraria, Centro de Estudos Agrarios, A, No. 1: (in French). Little, T. M. and Hills, F. J. (1975). Statistical methods in agricultural research. DAVIS, University of California. Mustata, G. (1992). Role of parasitoid complex in limiting the population of diamondback moth in Moldavia, Romania. In Diamondback moth and other crucifer pests: Proceedings of the second international workshop (ed. Talekar, N. S. and Griggs, T. D.) pp Shanhua, Taiwan: Asian Vegetable Research and Development Center. Talekar, N. S. and Yang, J. C. (1991). Characteristic of parasitism of diamondback moth by two larval parasites. Entomophaga. 36: Talekar, N. S. (1996a). Role of biological control in IPM of diamondback moth in Southeast Asia. Paper presented at the Workshop on Biological Control as a Cornerstone of IPM for Sustainable Agriculture in Southeast Asia, Serdang, Malaysia (in Press). Talekar, N. S. (1996b). Biological control of diamondback moth in Taiwan a review. Plant Protection Bulletin (Taiwan), (in press) Velasco, L. R. I. (1983). Field parasitism of Apanteles plutellae Kurdj (Braconidae: Hymenoptera) on the diamondback moth on cabbage. Philippine Entomologist 6: Wakisaka, S., Tsukuda, R. and Nakasuji, F. (1991). Life tables of diamondback moth, Plutella xylostella (L.) (Lepidoptera: Yponomeutidae) and effects of rainfall temperature and host plants on survival and reproduction. Japanese Journal of Applied Entomology and Zoology 35: Proceedings: The Management of Diamondback Moth and Other Crucifer Pests

7 Wu, K. C. (1968). Bionomic study of Plutella maculipennis Curtis. Journal of Taiwan Agricultural Research 17: Yang, J. C., Chu, Y. I. and Talekar, N. S. (1994). Studies on the characteristics of parasitism of Plutella xylostella (Lep.: Plutellidae) by a larval parasite Diadegma semiclausum (Hym.: Ichneumonidae). Entomophaga 39: Yaseen, M. (1978). The establishment of two parasites of the diamondback moth, Plutella xylostella (Lepidoptera, Plutellidae) in Trinidad, W. I. Entomophaga 23: Biologically-based technologies 13

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