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Homalodisca Coagulata Egg Masses: Economic Implications for Mass Rearing and Biological Control

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Homalodisca Coagulata Egg Masses: Economic Implications for Mass Rearing and Biological Control This article was originally published in a journal published by Elsevier, and the attached copy is provided by Elsevier for the author’s benefit and for the benefit of the author’s institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues that you know, and providing a copy to your institution’s administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier’s permissions site at: http://www.elsevier.com/locate/permissionusematerial Biological Control 39 (2006) 162–170 www.elsevier.com/locate/ybcon The eVect of intraspeciWc competition on progeny sex ratio in Gonatocerus spp. for Homalodisca coagulata egg masses: Economic implications for mass rearing and biological control N.A. Irvin ¤, M.S. Hoddle Department of Entomology, University of California, 3041 Watkins Drive, Riverside, CA 92521, USA Received 9 January 2006; accepted 16 May 2006 Available online 20 May 2006 Abstract The eVect of the number of simultaneously ovipositing females on progeny sex ratio of three Homalodisca coagulata egg parasitoids, Gonatocerus ashmeadi, G. triguttatus, and G. fasciatus was investigated in the laboratory. When one female Gonatocerus parasitoid was present, progeny production was strongly female biased producing »1 male:8 females, 1:14 and 1:9 for G. ashmeadi, G. triguttatus and G. fasciatus, respectively. Increasing the number of simultaneous ovipositing females from 1 to 3, signiWcantly increased percentage of male oVspring by up to 61% for all three Gonatocerus species. Overall percentage male oVspring ranged from 31–34%, and did not signiW- cantly diVer among species. Overall mean percentage parasitism by G. ashmeadi was up to 16% higher compared with G. triguttatus and G. fasciatus, whereas, average clutch size and mean progeny production was more than twofold higher for G. fasciatus when compared with G. ashmeadi and G. triguttatus. Additional laboratory trials with only G. ashmeadi demonstrated that when H. coagulata egg masses consisted of 2–13 eggs, 67–100% of females allocated one male oVspring per egg mass. When one egg mass was presented to Wve female G. ashmeadi searching concurrently, male oVspring was up to 22% higher compared with male oVspring from host eggs exposed to one female G. ashmeadi. Experiments indicated that allocation of male eggs was not inXuenced by constant female-to-female contact for 19 h prior to host exposure, and female G. ashmeadi did not alter allocation of male eggs when sibling and non-sibling conspeciWcs were con- testing host egg masses. The economic signiWcance for mass rearing is substantial. For G. ashmeadi at 2003 production costs and under optimal conditions, estimated cost per female was $1.42, under suboptimal conditions, cost per female increased to $4.51; a 218% increase in production cost per female parasitoid. © 2006 Elsevier Inc. All rights reserved. Keywords: Biological control; Gonatocerus ashmeadi; Gonatocerus fasciatus; Gonatocerus triguttatus; Homalodisca coagulata; IntraspeciWc competition; Local mate competition; Mass rearing; Sex ratio 1. Introduction et al., 2003), it readily vectors a xylem-dwelling bacterium, Xylella fastidiosa Wells et al. (Adlerz, 1980). DiVerent The glassy-winged sharpshooter, Homalodisca coagulata strains of this bacterium induce scorch-like disease in over (Say) (Hemiptera: Cicadellidae), established in California, 100 host plants including grapes, almonds, plums, peaches, USA in the late 1980s, and is creating serious new economic alfalfa, oleander and liquidambar (Purcell and Saunders, problems in both agricultural and urban landscapes. As this 1999; Costa et al., 2000; Hopkins and Purcell, 2002). In insect continues to spread throughout California, feeding 2001, the California Department for Food and Agriculture from xylem tissue of overAuthor's 100 known host plants ( Hoddle personal(CDFA) initiated a classical copy biological control program for H. coagulata to reduce densities of this pest and the associated spread of Xylella-related diseases. Mymarid egg * Corresponding author. Fax: +1 951 827 3086. parasitoids, Gonatocerus ashmeadi Girault, G. triguttatus E-mail address: [email protected] (N.A. Irvin). Girault, and G. fasciatus Girault have been imported, 1049-9644/$ - see front matter © 2006 Elsevier Inc. All rights reserved. doi:10.1016/j.biocontrol.2006.05.005 N.A. Irvin, M.S. Hoddle / Biological Control 39 (2006) 162–170 163 mass-reared and released in California as part of this bio- ognized sibling-conspeciWcs during oviposition. The second logical control program. G. ashmeadi has been resident in study examined the eVect of size of egg masses on progeny California since 1978 (Huber, 1988), while G. triguttatus sex ratio. It was hypothesized that Gonatocerus spp. would was introduced from Texas in 2001. In 2002, G. fasciatus increase male progeny production in response to increasing from Louisiana was moved through quarantine and cleared female density, as predicted by LMC. It was also hypothe- for release in California (CDFA, 2003). G. ashmeadi and sized according to LMC that when G. ashmeadi is presented G. triguttatus are solitary endoparasites while G. fasciatus is with one egg mass of varying size, one son would be allo- a gregarious endoparasitoid (Triapitsyn et al., 2003). cated to a speciWc number of female oVspring and this ratio Gonatocerus ashmeadi, G. triguttatus and G. fasciatus of male to female oVspring would be constant across host produce female biased sex ratios of up to 79, 77 and 96s, egg masses of varying sizes. respectively (Triapitsyn et al., 2003; Irvin and Hoddle, 2005a; Irvin and Hoddle, 2005b). All three parasitoid spe- 2. Materials and methods cies are haplodiploid and can manipulate oVspring sex ratio by storing sperm from a mating and controlling the access 2.1. Insect colonies of sperm to eggs at the time of oviposition (Flanders, 1965; Green et al., 1982). Local mate competition (LMC) (Hamil- Laboratory colonies of H. coagulata, G. ashmeadi, G. tri- ton, 1967) proposes that female parasitoids should always guttatus and G. fasciatus were maintained at the University attempt to maximize their reproductive Wtness so when a of California, at Riverside. Parasitoid colonies were held at single female visits a patch of hosts, her Wtness is propor- 26 § 2 °C and 30–40% RH under a L14:10D photoperiod tional to the number of fertilized daughters emigrating and reared on H. coagulata eggs laid on ‘Eureka’ lemon from the patch (Green et al., 1982; Hardy, 1992; Hardy leaves, a preferred lemon variety for H. coagulata oviposi- et al., 1998). Consequently, the foundress should oviposit tion and parasitoid foraging (Irvin and Hoddle, 2004; [see just enough haploid sons in each patch to insure her diploid Irvin et al., 2006 for plant maintenance details]). All experi- daughters are inseminated. This maximizes female Wtness, ments were conducted in the laboratory at 26 § 2°C and ensures suYcient males are present to mate with females, 30–40% RH under light intensity of 1.2 § 0.2 log lumens/m2 and reduces competition between brothers for sib-mating and L14:D10 photoperiod. with sisters (Luck et al., 2001). When the number of ovipositing females accessing a 2.2. Progeny sex ratio resulting from 30 H. coagulata eggs host patch increases, LMC predicts that an increasing investment in sons versus daughters will be advantageous Lemon leaves containing »30 H. coagulata eggs were because sons will distribute a greater number of genes to placed through holes drilled through the lid of a 130 ml females from unrelated lineages thereby increasing the plastic vial (40 dram Plastic Vial, Thornton Plastics, Salt Wtness of their own lineage (Grafen, 1984; Luck et al., 1999; Lake City, UT) Wlled with deionized water and 3 ml of anti- Luck et al., 2001). Consequently, this phenomenon of septic (Listerine Antiseptic Mouthwash, PWzer Inc., New increasing male production with increasing densities of ovi- York, NY) to prevent bacterial rot. Eggs were approxi- positing females can have detrimental consequences for mately 24–48 h of age (shown to be successfully utilized by mass rearing programs which aim to maximize the produc- all three Gonatocerus spp. [Irvin and Hoddle, 2005a]). A tion of females for colony maintenance and release as bio- second 130 ml plastic vial with ventilation (three 2 cm holes logical control agents (Waage, 1982; Waage et al., 1984; [one on the bottom, and one on each of two sides] covered Hall, 1993). This is particularly relevant for H. coagulata with mesh netting [80 m JelliV Corporation, Southport, which is extremely diYcult to economically mass rear, and CT]) was inverted and attached to the lid of the vial holding almost impossible to maintain in colony year round. the water and lemon leaves. A drop of honey-water (3:1 Together, these two factors make egg masses from this pest Natural uncooked honey, Wild Mountain Brand, Oakland, extremely valuable for parasitoid rearing. Therefore, identi- CA) was placed on the side of the inverted vial that covered fying and understanding factors inXuencing sex ratio of the test material and one newly emerged male and female Gonatocerus spp. is imperative for maintaining successful parasitoid (»24 h old) were introduced. This procedure was and eYcient mass rearing programs and for interpreting replicated approximately 15 times for each of Wve female parasitoid oVspring production from host patches under densities for each parasitoid species. Female densities tested Weld conditions (Waage et al., 1984; Waage, 1986). Here, we were 1, 2, 3, 4, and 5 G. ashmeadi, G. triguttatus, and G. fas- investigate the eVect ofAuthor's female density on progeny sex personalratio ciatus. For each female densitycopy tested, one male parasitoid of three H. coagulata egg parasitoids, G. ashmeadi, G. tri- of the same species was simultaneously present in test vials guttatus and G.
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