Prediction of the Life Cycle of the West Japan Type Yellow-Spotted Longicorn Beetle, Psacothea Hilaris (Coleoptera: Cerambycidae) by Numerical Simulations
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Appl. Entomol. Zool. 37 (4): 559–569 (2002) Prediction of the life cycle of the west Japan type yellow-spotted longicorn beetle, Psacothea hilaris (Coleoptera: Cerambycidae) by numerical simulations Yuya Watari, Takehiko Yamanaka, Wataru Asano and Yukio Ishikawa* Laboratory of Applied Entomology, Department of Agricultural and Environmental Biology, The University of Tokyo, Tokyo 113– 8657, Japan (Received 3 December 2001; Accepted 3 July 2002) Abstract A temporally structured model that enables simulation of the development of the west Japan type yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe), at different locations was developed. Life history parameter values in- corporated into the model were estimated by laboratory rearing experiments. To validate the present model, the devel- opment of eggs laid monthly from June 1 through November 1 was simulated under dynamic temperature and pho- toperiod conditions at Ayabe City. The individuals laid on June 1 did not enter diapause but emerged in early August of the same year. On the other hand, about 2/3 of the individuals laid on July 1, and all those laid on August 1 and September 1 entered diapause (or quiescence), and started to emerge in late May of the following year. Individuals laid on October 1 and November 1 overwintered as young larvae (1st–3rd stadia) and eggs, respectively, and the ma- jority of these emerged in late July–early August. Interestingly, the remaining individuals entered diapause in the 2nd year and emerged in June of the 3rd year. Analyses of these simulation results suggested that concentrated emergence of P. hilaris can occur twice in one year (in late May–early June and in late July–early August) at Ayabe, and this is fairly concordant with known adult prevalence at this location considering the long life-span of adults. It was also sug- gested that although P. hilaris at Ayabe has basically a univoltine life cycle with a facultative diapause, varying pro- portions of the population appear to have a bivoltine life cycle or a semivoltine life cycle depending on the meteoro- logical conditions of that year. The life cycle of P. hilaris is suggested to be very flexible and adaptive to environmen- tal fluctuations. Key words: Psacothea hilaris, seasonal prevalence, temporally structured model, bivoltine life cycle, semivoltine life cycle continuous stripes (Iba, 1980). INTRODUCTION These two types are also characterized by differ- The yellow-spotted longicorn beetle, Psacothea ent seasonal life cycles and photoperiodic responses hilaris (Pascoe), is widely distributed in eastern- (Iba et al., 1976; Sakakibara and Kawakami, 1992). most Asia. This species is known to have large ge- The west Japan type has its main adult prevalence ographic variation in morphology, and is classified peak in early summer, and a small second peak is into 13 subspecies (Kusama and Takakuwa, 1984). observed in autumn (Iba et al., 1976; Suzuki and One subspecies, P. hilaris hilaris, inhabits Honshu, Yoshii, 1980; Ishiishikawa, 1986). They are consid- Shikoku and Kyushu, three of the four main islands ered to enter diapause as mature larvae in response of Japan. Within this subspecies, two morphologi- to low temperatures and short daylength in autumn cal types, the “west Japan type” and “east Japan (Shintani et al., 1996b; Shintani and Ishikawa, type” have been recognized (Iba, 1980; Makihara, 1997a). On the other hand, the east Japan type does 1986). The west Japan type inhabits Kyushu, not form a sharp emergence peak, but an increase Shikoku and western parts of Honshu, and is char- in the adult population is usually found in autumn acterized by severed yellow stripes on the prono- (Iba, 1976; Iba et al., 1976; Sakakibara and Kawa- tum. In contrast, the east Japan type is found in the kami, 1992). They had been regarded to lack dia- eastern parts of Honshu, and is characterized by pause and overwinter at the egg stage (Makihara, * To whom correspondence should be addressed at: E-mail: [email protected] 559 560 Y. Watari et al. 1986; Sakakibara and Kawakami, 1992), however, tion Shintani and Ishikawa (1997a) have shown that 1/D5(T2T )/K (1) they also enter diapause but at temperatures lower 0 than for the west Japan type. where D is the duration of a developmental stage in P. hilaris is a serious pest of mulberry and fig days, T is the constant rearing temperature, K is the trees. Control of P. hilaris is difficult because lar- thermal constant for the stage, and T0 is the lower vae bore tunnels in the trunks and are thus invisible thermal threshold for development (developmental from the outside. In addition, extensive insecticide zero). treatment of mulberry trees is not practical since Except for the egg stage, K and T0 were esti- mulberry leaves are fed to the silkworm, Bombyx mated from the information obtained from the mori. P. hilaris damages host trees heavily when above rearing experiments using a linear regression they are larvae, particularly after the 3rd stadium of 1/D against T (see Results). K and T0 for the egg (Iba, 1993). Thus, prediction of the detailed age- stage were obtained from Sakakibara (1995), who structure is indispensable to determine the best determined these values for P. hilaris from various time to control P. hilaris. locations in Japan. The values for the Kochi City The objective of the present study was to con- population (west Japan type) were used in the pres- struct a temporally structured model for prediction ent simulations. of the time of adult emergence, and to compare the Critical daylength. Shintani and Ishikawa (1998) simulation results with previous findings on adult reported that the critical daylength (CDL) for dia- prevalence in the field. Since the larval photoperi- pause induction of the west Japan type P. hilaris odic responses of P. hilaris have mainly been falls between 13.5 and 14 h at 25°C. However, they investigated using the west Japan type population did not determine the CDL at other temperatures. (Shintani et al., 1996a; Shintani and Ishikawa, In the present simulation, CDL was set as 13.75 h 1997a, b, c, 1998), in the present study, we con- (the midpoint of 13.5 h and 14 h) regardless of the structed a model for the west Japan type P. hilaris temperature, and used as a parameter that switches incorporating its daily age-structure, larval and long-day type responses and short-day type re- pupal development, and local meteorological data. sponses. Namely, when the daylength of the day t (DLt) was greater than CDL, temperature depend- ent responses similar to those at 15L : 9D were MATERIALS AND METHODS # assumed to occur. Conversely, when DLt CDL, Parameter estimation responses similar to those at 12L : 12D were as- Rearing experiment. Adults of the west Japan sumed. type P. hilaris were collected from mulberry fields Meteorological parameter values incorporated in Ino Town (33.5°N, 133.4°E), Kochi Prefecture, into the simulation. The daylengths between Janu- and maintained in the laboratory at 25°C under a ary 1 and December 31 at Ayabe were calculated 15 h light–9 h dark photoregime (15L : 9D). Eggs by the sunrise and sunset times in 2001 down- were obtained as described by Shintani et al. loaded from the Hydrographic Department, Japan (1996a). Newly-hatched larvae were individually Coast Guard (http://www.jhd.go.jp/cue/KOHO/ placed in plastic Petri dishes (6 cm diam.31 cm) automail/sun_form4.htm). The daily maximum, with a piece of artificial diet (Silkmate 2S®, Nihon mean and minimum air temperatures averaged for Nosan K.K., Yokohama, Japan). Sixty to 100 lar- eight years between 1960 and 1967 at Kyoto mete- vae each were subjected to ten combinations of two orological station, which is the station nearest to photoregimes (12L : 12D and 15L : 9D)3five tem- Ayabe City, were obtained from the Monthly Re- peratures (20.0, 22.5, 25.0, 27.5 and 30.060.1°C). port of the Japan Meteorological Agency. Molting, pupation and adult eclosion were checked Model construction daily for 180 d from the start of the experiment. Framework. The daily-structured model was de- Thermal constants and developmental thresholds veloped using Microsoft Developer Studio (version for each stage. It is well-known that the growth rate 4.0, Microsoft®) with C11 programming lan- of insects is proportionate to temperature within an guage. The model has 365 time steps for a one- optimum range, and can be expressed by the equa- year season, and each time step (t) corresponds to Life Cycle of West Japan Type P. hilaris 561 Fig. 1. The schematic flowchart of procedures in the simulation model. Arrows indicate the flows of the processes in one time step (one day). each day between January 1 and December 31. The riod and the mean temperature as described below. calculations were started on the day of oviposition Individuals were assumed not to suffer from any (D0) and continued until all individuals emerged as mortality; all survived until the end of the simula- adults. During the simulation period, 100 individu- tion. These procedures are shown in Fig. 1 as a # , als (YSLi :0 i 100) grew uniformly at a daily rate schematic flow chart. determined by the maximum and minimum tem- Growing process dependent on the temperature. peratures of the day (see the next section) until the The stage and mode of the i-th individual were des- 5 end of the 3rd stadium. Thereafter, the individuals’ ignated as Si ( egg, 1st, 4th, ...9th or pupa) and 5 destiny (molting, pupation or diapause) bifurcated Mi ( normal, pupation or diapause), respectively. at the end of each stadium based on the photope- The cumulative heat unit (degree-days) was used as 562 Y.