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Thermal Requirement for Development of Carpophilus Marginellus

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Thermal Requirement for Development of Carpophilus Marginellus Appl. Entomol. Zool. 43 (2): 281–285 (2008) http://odokon.org/ Thermal requirement for development of Carpophilus marginellus (Coleoptera: Nitidulidae), a potential pollinator of cherimoya and atemoya trees (Magnoliales: Annonaceae) Morio TSUKADA,1,* Daisuke TANAKA1 and Hirokazu HIGUCHI2 1 Insect Ecology Laboratory, Faculty of Bioresources, Mie University; Mie 514–8507, Japan 2 Laboratory of Tropical Agriculture, Faculty of Agriculture, Kyoto University; Kyoto 606–8502, Japan (Received 22 February 2007; Accepted 21 January 2008) Abstract Carpophilus marginellus is a candidate pollinator of the subtropical orchard trees cherimoya and atemoya. We investi- gated the developmental period and adult size of this species at constant temperatures of 18, 20, 25, and 30°C as basic information for mass rearing and release in greenhouses as a pollinator. The photoperiod was 16-h light/8-h dark at all temperatures. The developmental period was shorter and the adult size was smaller at higher temperatures. Mortality was lowest at 25°C and highest at 30°C. The thermal threshold (developmental zero) ranged from 11.9 (female larvae) to 15.6°C (female pupae) among developmental stages and sexes. The thermal constant (total effective temperature) of the period from egg to adult emergence was ca. 292 degree-days. Key words: Degree-days; developmental zero; adult size; Annona; beetle pollination Such facilities are favorable for the use of mass- INTRODUCTION released insect pollinators, since the greenhouse The cherimoya, Annona cherimola Mill. (Mag- can prevent the insects from escaping. We have noliales: Annonaceae), is a subtropical orchard been studying visitors to cherimoya flowers in tree. Although it is self-compatible, autogamy is Japanese orchards for the purpose of finding suit- generally impossible because of protogynous di- able pollinators during the flowering season chogamy (Sanewski, 1991). This is a relatively (Tsukada et al., 2005b). Annona spp. floral differ- archaic plant that does not use bees as pollinators. entiation occurs year-round in their natural distri- Thus, hand pollination is used to obtain an ade- bution area. However, in Honshu, pruning in early quate fruit set, but this takes time and money spring results in full-blooming around June. We (Gazit et al., 1982; Richardson and Anderson, found several individuals of Carpophilus marginel- 1996). Several studies have been conducted to lus Motschulsky (Coleoptera: Nitidulidae) on the identify suitable pollinators of cherimoya and its flowers, among other species such as Mimemodes relative, atemoya (A. cherimolaϫsquamosa), to monstrosus (Reitter) (Rhizophagidae) and Hapton- reduce orchard labor. Nitidulid beetles are the main cus ocularis (Fairmaire) (Nitidulidae) during the visitors of Annona spp. flowers world widely flowering season. (Gazit et al., 1982; George et al., 1989; Nagel et We have already reported the developmental al., 1989; Nadel and Peña, 1994), and secondary characteristics of H. ocularis as a candidate polli- visitors include staphylinid beetles and Orius spp. nator (Tsukada et al., 2005a). However, high mor- (Hemiptera: Anthocoridae) (Caleca et al., 1996, tality at high temperatures limits its use in green- 1998; Palmeri and Longo, 1997). houses during summer (Higuchi et al., unpub- These fruit trees were introduced to the Japanese lished). This prompted us to investigate another main island, Honshu, two decades ago. They are candidate pollinator. Among the flower visitors in cultivated in greenhouses to avoid frost damage. Japan, the number of C. marginellus was second *To whom correspondence should be addressed at: E-mail: [email protected] DOI: 10.1303/aez.2008.281 281 282 M. TSUKADA et al. largest, after M. monstrosus (Tsukada et al., 2005b). moya in an experimental orchard of Mie Univer- Both males and females enter the female-stage sity, central Japan. They were reared in plastic con- flowers and leave them during the male stage with tainers measuring 26 cmϫ18 cmϫ8 cm (height). much pollen on their bodies (unpublished data). Three holes of 4-cm diameter were made in the lid Therefore, we assume that they pollinate the plant. of each container and covered with a fine gauze, This sap beetle has a body length of ca. 3 mm which allowed sufficient ventilation but prevented (Hisamatsu, 1985), and is distributed widely in insects from escaping. Cut pineapple was periodi- warm regions from East Africa to Japan (Kirejt- cally supplied as food, and the bottom of the con- shuk, 1998). Females lay their eggs on rotting tainer was lined with autoclave-sterilized soil for fruits such as orange and pineapple. Larvae de- pupation. Water was sprayed as needed on the soil. velop there, and pass through three instars before The beetles were easily reared at 25°C in a 16-h pupation. Mature last-instar larvae wander about light/8-h dark (16L8D) photoperiod. before entering the soil, where they pupate. Larvae Development from egg to adult. The procedure also develop on ripened Annona fruits. However, for the rearing of insects was similar to that used Annona fruits usually ripen after harvest, so for H. ocularis (Tsukada et al., 2005a). The fe- C. marginellus is not likely to damage the fruit in males laid their eggs into the pineapple, but be- orchards. Also, even though some flower visitors, cause finding eggs there was extremely difficult, including nitidulid beetles, are known to injure we used the following procedure to obtain eggs. plant ovaries, which causes aesthetic damage to First, 52 pairs of male and female were prepared. some orchard fruits, this seems not to occur on the Each was allowed to oviposit into a cut pineapple cherimoya (personal observation). Thus, the use of in a 30-ml vial in a chamber kept at 18, 20, 25, or this species as a pollinator by mass release in 30°C under a 16L8D photoperiod. The opening of greenhouses is feasible. On the other hand, C. mar- the vial was plugged by cotton to allow moderate ginellus, as well as other Carpophilus species, can ventilation. The insects were then removed within a be pests of dried fruits and cereals, but adequate day and placed into a new vial with another piece pest management has not been established, because of pineapple for the next oviposition. This proce- of the lack of published data on their basic biology. dure was repeated until sufficient larvae were ob- To investigate the use of beetles as pollinators of tained. Each vial with the pineapple piece was kept the cherimoya, especially by mass rearing and at the same temperature, and the pieces were exam- release in greenhouse orchards, researchers must ined every day. When larvae were found, they were know their developmental characteristics. Among isolated and each was gently transferred to a 10-ml many abiotic environmental factors, circumstantial vial with a ca. 5 g pineapple piece as food and a temperature is one of the most crucial factors that piece of wet tissue paper as a pupation site. Usu- regulate the development of insects. Except at ally, there was no need to add another peace of extremely high or low temperatures, insects de- pineapple, but we added one when the food ap- velop faster and attain smaller adult size at higher peared insufficient. The tissue paper was ca. 0.4 g temperatures (Atkinson, 1994). Data of develop- in weight and contained ca. 1.2 ml of water, a suit- mental rate and adult size offers basic information able volume for the pupation of C. marginellus for the mass rearing of insects. In this study, we (unpublished data). The vials were examined every experimentally elucidated the thermal requirements day, and pupation date and adult emergence date and adult size of C. marginellus. Our results offer were recorded. The dates of oviposition and ap- basic information both for the use of C. marginel- pearance of new larvae (regarded as egg hatch) lus as a pollinator for cherimoya and other Annona were also recorded. The sex of each individual was plants, and for management of it as a pest of dried determined after adult eclosion by observation of fruits. the tip of the abdomen under a binocular micro- scope. Adult size. The emerged adults were individu- MATERIALS AND METHODS ally dried, and head width (distance between ante- Insect stock culture. The insects originated rior ends of the compound eye) was measured from feral individuals caught in the flowers of ate- using a micrometer under a binocular microscope. Development of Carpophilus marginellus 283 Table1.Developmental period (days) of C. marginellus at four temperatures. MeanϮSD Temp. Sex (n) Egg Larva Pupa Total 18°C ? (44) 12.41Ϯ1.69 25.73Ϯ1.78 22.20Ϯ2.08 60.07Ϯ4.04 / (42) 12.76Ϯ1.49 25.36Ϯ1.48 22.00Ϯ2.54 59.64Ϯ3.99 20°C ? (35) 9.26Ϯ1.48 22.06Ϯ1.92 17.94Ϯ1.63 49.83Ϯ3.33 / (36) 8.78Ϯ1.17 21.81Ϯ2.07 18.75Ϯ2.59 49.28Ϯ3.78 25°C ? (46) 5.24Ϯ0.97 12.96Ϯ1.35 7.96Ϯ1.03 26.22Ϯ1.95 / (40) 5.20Ϯ0.91 12.75Ϯ1.33 7.68Ϯ1.10 25.63Ϯ1.90 30°C ? (27) 4.44Ϯ0.80 8.89Ϯ1.40 5.07Ϯ1.07 18.41Ϯ1.53 / (33) 4.48Ϯ0.94 9.15Ϯ1.28 4.79Ϯ1.05 18.45Ϯ1.62 Statistical procedures. The parameters of ther- Table2.Total effective temperature (K; degree-days) and mal requirements (i.e., thermal constant [K] and the developmental zero (T0) of C. marginellus. The data at 30°C were omitted from calculation of parameters of egg stage thermal threshold [T0]) were obtained from the regression of 1/D against T, where D is days Stage Sex r2 a KT required to complete a stage and T is temperature 0 (°C). These regressions were estimated from the Egg ? 0.8132 60.24 13.16 raw data, not from the means of each temperature.
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