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Host-Substrate Preference of Theocolax Elegans (Westwood

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Host-Substrate Preference of Theocolax Elegans (Westwood Accepted Manuscript Host-substrate preference of Theocolax elegans (Westwood) (Hymenoptera: Pteromalidae), a larval parasitoid of the maize weevil, Sitophilus zeamais (Motschulsky) (Coleoptera: Curculionidae) Saruta Sitthichaiyakul, Weerawan Amornsak PII: S2452-316X(17)30089-3 DOI: 10.1016/j.anres.2016.09.003 Reference: ANRES 84 To appear in: Agriculture and Natural Resources Received Date: 29 July 2016 Accepted Date: 7 September 2016 Please cite this article as: Sitthichaiyakul S, Amornsak W, Host-substrate preference of Theocolax elegans (Westwood) (Hymenoptera: Pteromalidae), a larval parasitoid of the maize weevil, Sitophilus zeamais (Motschulsky) (Coleoptera: Curculionidae), Agriculture and Natural Resources (2017), doi: 10.1016/j.anres.2016.09.003. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. 1 ACCEPTED MANUSCRIPT 1 Agriculture and Natural Resources. 2016. 51(1): xx–xx. 2 Agr. Nat. Resour. 2016. 51(1): xx–xx. 3 4 Host-substrate preference of Theocolax elegans (Westwood) (Hymenoptera: 5 Pteromalidae), a larval parasitoid of the maize weevil, Sitophilus zeamais 6 (Motschulsky) (Coleoptera: Curculionidae) 7 8 Saruta Sitthichaiyakul a, b,† and Weerawan Amornsak a,* 9 10 a Department of Entomology, Faculty of Agriculture, Kasetsart University, Bangkok 11 10900, Thailand. 12 b Post-harvest and Processing Research and Development Division, Department of 13 Agriculture, Bangkok 10900, Thailand. 14 15 Received 29 July 2016 16 Accepted 7 September 2016 17 18 Keywords: MANUSCRIPT 19 Biological control 20 Host substrates 21 Sitophilus zeamais 22 Stored-product insects 23 Theocolax elegans 24 25 *Corresponding author. 26 E-mail: [email protected] 27 †Co-first author. 28 E-mail: [email protected] 29 30 31 32 33 34 2 ACCEPTED MANUSCRIPT 1 Abstract 2 3 The solitary parasitoid Theocolax elegans (Westwood) (Hymenoptera: 4 Pteromalidae) was investigated attacking larvae of the maize weevil, Sitophilus 5 zeamais (Motschulsky) (Coleoptera: Curculionidae) under laboratory conditions. 6 Theocolax elegans parasitoids were mass reared on 21-day-old S. zeamais fed with 7 different host substrates consisting of brown rice, maize, sorghum and wheat. The 8 developmental time of S. zeamais was observed. The widest head capsule was 9 recorded from S. zeamais developing in brown rice grain kernels. The head capsule 10 width was used to determine the age of the larval instars. The sex ratio of T. elegans 11 progeny emerging from brown rice was the same in the choice and no-choice tests 12 (1.8:1.0 and 1.8:1.0, respectively). Female parasitoids preferred to oviposit on S. 13 zeamais developed in brown rice grain kernels in both tests. The number of parasitoid 14 progeny emerging from different host substrates was different in the choice and no- 15 choice tests. The progeny of T. elegans females and males were fully winged, short 16 winged and wingless. 17 18 Introduction MANUSCRIPT 19 20 Stored-product insect pests infest grain stores around the world, with serious 21 damage to stored grains being caused by stored-product insect pests such as the rice 22 weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae) (Toews et al ., 2007), red 23 flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae) (Campbell 24 and Hagstrum, 2002), cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: 25 Anobidae) (Timokhov and Gokhman, 2003), and cowpea weevil, Callosobruchus 26 maculatus (F.) (Coleoptera: Bruchidae) (Ghimire and Phillips, 2007). These pests 27 cause economic loss in terms of the quality and quantity of the products. Biological 28 control is aACCEPTED method used to control stored-product insect pests and the potential 29 benefit of natural enemies such as parasitoids has been acknowledged for many years 30 (Bellows, 1985; Shin et al ., 1994; Adarkwah et al ., 2014). However, insecticides 31 remain the primary tool for controlling stored product insect pests, and consequently, 32 resistance to fumigation and contact insecticides has been reported with some stored 33 product insects (Boyer et al ., 2012; Kang et al ., 2013). 3 ACCEPTED MANUSCRIPT 1 Pteromalid parasitoids are important biological control agents (Howard and 2 Liang, 1993) and occur naturally in stored grain (Williams and Floyd, 1971). Species 3 include Anisopteromalus calandrae (Howard), Lariophagus distinguendus (Förster) 4 (Shin et al ., 1994) and Pteromalus cerealellae (Ashmead) (Wen et al ., 1995). 5 Theocolax elegans (Westwood) is a pteromalid ectoparasitoid used to suppress the 6 larval stage of several stored-product insect pests (Wen and Brower, 1995). These 7 beneficial parasitoids have been shown to attack coleopteran and lepidopteran insect 8 pests (Flinn et al ., 1994). They have been used as a biological control agent in stored 9 grain (Gordh, 1979; Germinara et al ., 2009). Flinn and Hagstrum (2001) reported that 10 augmentative releases of T. elegans reduced damage from the lesser grain borer, 11 Rhyzopertha dominica (F.) (Coleoptera: Bostrichidae). Investigations of T. elegans 12 larvae revealed survival and the preference of the parasite for the fourth and pupal 13 stages of the host (Sharifi, 1972; Flinn and Hagstrum, 2001). The effect of 14 temperature was reported to be a functional response in the reproduction of the 15 progeny of the parasitoid T. elegans (Flinn, 1998; Toews et al ., 1998; Toews et al ., 16 2001; Flinn and Hagstrum, 2002). The successful parasitism of the female parasitoid 17 depends on many factors including finding a target host (Steidle and Schöller, 2002), 18 acceptance of the target host and the developmentMANUSCRIPT of target host size (Bergeijk et al ., 19 1989; Bourchier et al ., 1993). Parasitoids acquire nutrients during their larval 20 development (Strand and Casas, 2008). Host preference and host suitability have been 21 studied in pteromalid parasitoids using different host species in stored grain 22 (Timokhov and Gokhman, 2003; Grimire and Phillips, 2007). Host quality was 23 related to host size and affected parasitoid progeny allocation (Cônsoli and Vinson, 24 2012). However, little information on T. elegans is available compared with other 25 pteromalid parasitoids. The current study provided basic information on host substrate 26 preferences by T. elegans on larval S. zeamais . 27 28 Materials andACCEPTED Methods 29 Insects host and parasitoid 30 Sitophilus zeamais was taken from silos in Amnatcharoen province, Thailand 31 during January 2013. Theocolax elegans was taken during 2011 from silos in 32 Petchaburi province, Thailand. These hosts and parasitoids were maintained at the 33 Post-harvest and Processing Research and Development Division, Department of 34 Agriculture, Chatuchak, Bangkok, Thailand. Both insects were mass reared at the 4 ACCEPTED MANUSCRIPT 1 National Biological Control Research Center, Kasetsart University, Bangkhen 2 Campus, Bangkok, Thailand under laboratory conditions (24–26°C, 50–60% relative 3 humidity, 12 h light:12 h dark as the natural photoperiod). 4 Mass rearing 5 The host species ( S. zeamais ) was mass reared on brown rice ( Oryza sativa L. 6 (Poaceae)), maize ( Zea mays L. (Poaceae)), sorghum ( Sorghum bicolar (L.) Moench 7 (Poaceae)) and wheat ( Triticum aestivum L. (Poaceae)) using glass containers holding 8 50 g each of brown rice, maize, sorghum and wheat. One hundred unsexed adults of 9 S. zeamais were placed in each container. Each glass container (5.5 cm diameter, 15 10 cm tall) was covered with a filter paper. The adults of host species oviposited for 5 d 11 and were then removed. The bottle was then maintained under laboratory conditions 12 until the larvae were used for other trials. 13 Theocolax elegans was mass reared on S. zeamais feeding on brown rice. Ten 14 pairs (females and males) of T. elegans were released when the larvae of S. zeamais 15 were aged 21 d in a glass bottle container (5.5 cm diameter, 15 tall cm) with a filter 16 paper cover. Theocolax elegans were allowed to parasitize the hosts. 17 18 Head capsule size of Sitophilus zeamais MANUSCRIPT 19 Sitophilus zeamais was mass reared on brown rice, maize, sorghum and wheat 20 and then placed individually with 5 g of plant substrate in test tubes. Five sexed pairs 21 were determined based on the surface, size and shape of snout characteristics (Tolpo 22 and Morrison, 1965; Dobie et al ., 1984; Throne and Eubanks, 2002). The individual 23 females were released on substrate to lay eggs for 24 h and covered with filter paper. 24 After an additional 21 d, S. zeamais larvae were removed from the grain kernel and 25 the width of larval head capsules was measured under a compound microscope 26 (Olympus BH-2 BHS Research Microscope; Olympus Corp.; Tokyo, Japan). A visual 27 record was made using the software program Ulead VideoStudio SE DVD (©2007, 28 InterVideo DigitalACCEPTED Technology Corp; Freemont, CA, USA). The measurement of head 29 capsule size was under taken using the software program Image-Pro PLUS (version 30 6.0.0.260, Media Cybernetics Inc.; Rockville, MD, USA). 31 32 Host substrates 33 A glass cylinder (12.5 cm diameter, 12.5 cm height) was divided into quarters 34 using four acrylic partitions (Fig. 1). For the choice experiments, 50 g of infested 5 ACCEPTED MANUSCRIPT 1 brown rice, maize, sorghum and wheat with 21-day-old S. zeamais were placed in 2 each quadrant. An acrylic circle covered with a hole enabled parasitoids to contact the 3 hosts. Theocolax elegans female parasitoids mated and were fed with honey for 24 h 4 before being released in the center of the quadrants. The experiment was replicated 20 5 times. 6 In the no-choice experiments, glass bottles (5.5 cm diameter, 15 cm height) 7 containing 50 g of infested host substrates (brown rice, maize, sorghum or wheat), 8 separately were used as treatments.
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