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Lymantria Xylina) Eggs ൢ֙ـं (ᖪ 24: 43-52 (2004ٿέ៉ Formosan Entomol. 24: 43-52 (2004) Diapause Termination in Casuarina Moth (Lymantria xylina) Eggs Shaw-Yhi Hwang*, Chao-Chih Jeng, Tse-Chi Shen, Yi-Shan Shae and Chen-Shawn Liu Department of Entomology, National Chung Hsing University, Taichung 402, Taiwan ABSTRACT The casuarina moth, Lymantria xylina Swinhoe, is an important pest of hardwood and fruit trees. Overwintering casuarina moths enter an obligatory diapause as a pharate first instar larva. We assessed the relationship between chilling duration and temperature in terminating the diapause stage of casuarina moth eggs. Eggs of the casuarina moth were exposed to a variety of artificial chilling conditions that differed in duration and temperature. Results indicated that diapausing eggs require exposure to cold temperatures (9~15°C) followed by warm temperatures (27°C) for successful emergence in the laboratory. Eggs exposed to longer chilling periods required shorter incubation times to emerge than eggs exposed to shorter chilling periods. Regression analyses revealed a negative relationship between emer- gence time and chilling duration. The use of a proper chilling temperature and duration can effectively shorten diapause of casuarina moth eggs and make the routine rearing of casuarina moth populations possible. Key words: Lymantria xylina, egg diapause, chilling treatment Introduction fruit trees and other hardwoods by L. xylina indicate the real threat posed by The casuarina moth, Lymantria xylina this moth. Swinhoe (Lepidoptera: Lymantriidae), is Shortly after emergence and mating a major pest of casuarina (Casuarina in the summer, females of the casuarina equisetifolia) and acacia (Acacia confusa) moth lay a single egg mass consisting of forests, natural areas, and fruit tree 100~1000 eggs, which they cover with orchards in Taiwan and the eastern coast hairs from their abdomens (Shen et al., of mainland China (Chao et al., 1996). 2003). The eggs then enter an obligatory The number of recorded host plants for diapause as a pharate first instar larva, this moth includes and probably exceeds prior to the complete consumption of the 69 species of trees and shrubs, belonging extra embryonic yolk (Shen et al., 2003). to 29 families (Chang and Weng, 1985; The diapause mechanism and how this Chao et al., 1996). Current infestations of stage of development is regulated for the *Correspondence address e-mail: [email protected]! Diapause Termination in Casuarina Moth (Lymantria xylina) Eggs 43 casuarina moth, however, are poorly useful in effectively shortening the dia- understood. The casuarina moth is closely pause stage of the casuarina moth and related to the gypsy moth, L. dispar, making the routine rearing of casuarina which is a very serious forest pest in moth populations possible. North America. Like the casuarina moth, the gypsy moth also undergoes diapause Materials and Methods as a pharate first-instar, and its eggs hatch in April after an 8~9-month Casuarina moth eggs dormancy (Williams et al., 1990; Lee and Casuarina moth egg masses were Denlinger, 1996, 1997; Gray et al., 2001). collected from two heavily infested areas Past research has revealed that gypsy in central Taiwan in early July 2002. moth oviposition occurs in early to mid These infested areas were located at summer when temperatures are near Mingchien, Nantoa County (23º49.1'N, yearly highs, and the developing embryos 120º39.3'E) and Erhshui, Changhua complete the prediapause phase in County (23º50.0'N, 120º36.4'E). Vege- approximately 16 days at 25°C (Gray et tation was composed primarily of Acacia al., 1991). Well-developed gypsy moth confusa and Sterculia foetida in the first embryos then enter a diapause phase area and Euphoria longana in the second that typically lasts 8~9 months (Gray et area. About 100 egg masses were collec- al., 2001). Research has also indicated ted from each site (Shen et al., 2003). that diapause is more rapidly terminated In the laboratory, egg masses were with exposure to low temperatures first surface-sterilized by soaking in 0.1% (Masaki, 1956; Pantyukhov, 1964; Giese sodium hypochlorite with 1% Tween 80 and Cittadino, 1977). After sufficient ex- solution for 10 min., then rinsing under posure to low temperatures to terminate running tap water for 5 min. The egg diapause, embryos will hatch within 11~ masses were then air-dried. For the 18 days at 25°C (Gray et al., 1995). In assessment of individual eggs, egg masses summary, gypsy moth egg phenology and were first broken up, and the hairs were hatching have been well studied and are removed by rubbing the masses against generally understood, and this infor- sticky tape. The dehaired eggs from dif- mation has aided gypsy moth manage- ferent egg masses were pooled and placed ment programs (Gray et al., 2001). into Petri dishes (500 eggs per dish). To As with any other insect manage- ensure that all eggs had entered diapause, ment program, development of appro- the dehaired eggs were placed in an priate artificial rearing methods for the environmental chamber at 25°C with a casuarina moth in the laboratory depends 12:12 (L:D) photoperiod for 10 days (Gray on sufficient knowledge of its life cycle et al., 1991, 1995, 2001). After this treat- and basic developmental biology. The ment, 50 diapausing eggs were placed objective of this study was to investigate into a 1.5-ml Eppendrof vial and capped the relationship between chilling duration with cotton; vials were then assigned to and temperature in terminating the dia- an experimental treatment. pause stage of casuarina moth eggs. Eggs of the casuarina moth were exposed to a Experimental treatments variety of artificial wintering treatments Upon diapause initiation, five vials that differed in duration and temperature. (50 eggs per vial) were randomly assigned We were interested in determining how to each of 165 rearing regimes. Each these variables interact to influence regime was composed of an exposure to hatching rates and emergence timing. We an experimental chilling treatment. Chil- believe that this information will be ling treatments combined 33 durations ௐ˘ഇסᖪௐ˟˩αٿέ៉ 44 (60, 65, 70, 75, …, 215, and 220 d) and and 15ºC treatments were about 2~3%, five constant temperatures (6, 9, 12, 15, 4~6%, and 7~9%, respectively. Correlation and 27ºC). All treatments were subjected and linear regression analyses of average to a 12:12-h (L:D) regime. hatching rates versus chilling durations Upon completion of each chilling for all different temperature treatments treatment, five vials of eggs (50 eggs per revealed weak relationships (Fig. 1). There- vial) from each chilling treatment were fore, egg hatching rates of the casuarina removed and placed at a constant 27ºC in moth were not correlated with the cold a Percival growth chamber with a 12: exposure periods. 12-h L:D photoperiod. Egg hatching was checked for each vial every 2 days for 60 Effects of chilling duration and days following treatment. Means and temperature on emergence times standard errors of the hatching rate and There was a significant difference in the average emergence time were cal- the average emergence times of casuarina culated for each treatment. moth eggs among the different tempera- ture treatments (Table 2). At the 6ºC Statistical analysis treatment, almost all eggs failed to hatch The egg hatching rates and the aver- through the exposure period. For the age emergence times among different other three chilling treatments (9, 12, experimental treatments were analyzed and 15ºC), average emergence times varied by analysis of variance (ANOVA; PROC significantly among different exposure GLM; SAS Institute, 1998), followed by periods (Table 2). For these three tem- comparisons of means by the least-signi- perature treatments, the average emerg- ficant difference (LSD) test. Linear ence time decreased with increasing ex- regression analyses were used to relate posure duration. Correlation and linear average hatching rates and average regression analyses of average emergence emergence times with the durations of time versus exposure duration revealed different chilling treatments (PROC REG; negative relationships (Fig. 2). The stron- SAS Institute, 1998). gest and most negative relationship was found at the 15ºC treatment between the Results average emergence time and the exposure duration (r2 = 0.93, p < 0.001) (Fig. 2). Effects of chilling duration and Overall, our results suggest that by temperature on hatching rates observing chilling duration, the emerg- There were significant differences in ence time can be closely predicted for the average hatching rates of casuarina different temperature treatments (r2 > moth eggs among the different chilling 0.77, p < 0.001). treatments (Table 1). Generally, average hatching rates were very low for all of Discussion the experimental treatments. No eggs hatched from the 27ºC treatment. At the Our results indicated that diapausing 6ºC treatment, almost all eggs failed to casuarina moth eggs require exposure to hatch through the exposure period. Aver- cold temperatures followed by warm tem- age hatching rates varied significantly peratures for successful emergence in the among different exposure periods for the laboratory. This study also clearly demon- other three temperature treatments (Table strated that eggs exposed to longer 1). Average hatching rates increased with chilling periods require a shorter warm increasing temperature treatments, and temperature incubation for emergence the average hatching rates for the 9,
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