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Reproductive Biology and Functional Response of Dineulophus Journal of Insect Science: Vol. 12 | Article 153 Savino et al. Reproductive biology and functional response of Dineulophus phtorimaeae, a natural enemy of the tomato moth, Tuta Downloaded from https://academic.oup.com/jinsectscience/article-abstract/12/1/153/892634 by guest on 27 August 2019 absoluta Vivina Savino,1,2a* Carlos E. Coviella,1b and María G. Luna2c 1Programa de Ecología Terrestre, Departamento de Ciencias Básicas e Instituto de Ecología y Desarrollo Sustentable (INEDES). Universidad Nacional de Luján, Argentina. Rutas 5 y 7, Luján (6700), Argentina 2Centro de Estudios Parasitológicos y de Vectores (CEPAVE –CCT La Plata- CONICET-UNLP), Calle 2 Nro. 584 (1900) La Plata, Argentina Abstract The tomato moth, Tuta absoluta (Lepidoptera: Gelechiidae), is a major pest in South America and is at present an important invasive species in the Mediterranean Basin. The larval stadium mines leaves, stems, and fruits, and chemical control is the most used control method in both its original range and the invaded distribution regions. Since current T. absoluta control strategies seem limited, biological control is a prominent tool to be applied abroad. The naturally occurring larval ectoparasitoid in Argentina and Chile Dineulophus phtorimaeae (Hymenoptera: Eulophidae) has been reported to have potential biocontrol efficiency. In this study, the ovigeny strategy of D. phtorimaeae was analyzed throughout the adult female lifetime, and the functional response of females offered a range of 2–15 T. absoluta larvae was measured over a 48-hour period. Mean D. phtorimaeae egg load was 4.15 eggs, and egg production resulted in extremely synovigenic behavior. Meanwhile, a decreasing number of eggs, due to resorption, was found. Proportions of attacked (host-fed and/or parasitized) and only host-fed hosts by the ectoparasitoid were density independent for the tested host range, exhibiting a type I functional response to T. absoluta, with an attack rate of 0.20 host larvae. Meanings of this reproductive strategy in evolutionary time as well as the consequences for augmentative biological control programs are discussed. Keywords: host-feeding, ovigeny, parasitoid, type I functional response Correspondence: a [email protected], b [email protected], c [email protected], *Corresponding author Editor: Michael Strand was editor of this paper. Received: 31 October 2011 Accepted: 3 October 2012 Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 12, Number 153 Cite this paper as: Savino V, Coviella CE, Luna MG. 2012. Reproductive biology and functional response of Dineulophus phtorimaeae, a natural enemy of the tomato moth, Tuta absoluta. Journal of Insect Science 12:153. Available online: http://www.insectscience.org/12.153 Journal of Insect Science | www.insectscience.org 1 Journal of Insect Science: Vol. 12 | Article 153 Savino et al. Introduction García et al. 2005; Polack et al. 2002; Mollá et al. 2011). Since T. absoluta has a widespread The moth Tuta absoluta (Meyrick) distribution, the current control techniques (Lepidoptera: Gelechiidae), a pest of tomato, seem to be limited. Particularly, control by Solanum lycopersicum L. (Solanales: insecticides might be inefficient, as it is Solanaceae), causes significant economic loss complicated by the appearance of resistant to tomato cropping in many countries in South populations and, even more importantly, America (Botto et al. 1999; Siqueira et al. produces adverse effects to human health and Downloaded from https://academic.oup.com/jinsectscience/article-abstract/12/1/153/892634 by guest on 27 August 2019 2000; Magalhaes et al. 2001). Although this is to the environment (Lietti et al. 2005; Bielza an endemic Neotropical pest, it has acquired a 2010). As a feasible phytosanitary measure, wider geographical distribution after its biological control within an integrated pest unintended introduction in other tomato management program is a challenging but production regions, such as southern Europe potentially very beneficial tactic to develop. and northern Africa, during the last decade (EPPO Bulletin 2005; Urbaneja et al. 2007; Among a list of more than 50 species of T. Netherlands Plant Protection Service 2009; absoluta entomophagous insects reported Speranza et al. 2009; Desneux et al. 2010). In elsewhere (García Roa 1989; Consoli et al. its new regions, T. absoluta has spread 1998; Miranda et al. 1998; Faria et al. 2000; extremely fast, becoming a potential threat to Luna et al. 2010, 2011; Desneux et al. 2010; the world tomato production (Desneux et al. Medeiros et al. 2011), some of them have 2011). been pointed out as potential biocontrol agents by means of augmentative releases (Pratissoli T. absoluta damage is caused by the feeding et al. 1998; Giustolin et al. 2001; Torres et al. activity of larvae. All larval stadia feed on the 2002; Luna et al. 2007; Faria et al. 2008; mesophyll of the leaf and produce mines and Sánchez et al. 2009; Luna et al. 2010; Mollá et galleries that reduce the photosynthetic al. 2011). However, efficient control has not capacity of the plant. At high density levels, it been demonstrated yet, with basic biological can affect mature fruits and stems, and research still needed if a successful integrated indirectly generate an entrance way for pest management program is to be developed pathogens, thus severely degrading crop for this pest. commercial value. The larval ectoparasitoid Dineulophus Several strategies are commonly used to phtorimaeae de Santis (Hymenoptera: control T. absoluta in its original and invaded Eulophidae) is a species native to Argentina distribution regions: pest monitoring, and Chile and has been the subject of agricultural practices, chemical control, laboratory and field studies in order to gain management by means of semiochemicals knowledge on its systematics (de Santis 1983) (mating disruption and mass trapping), and to and basic biological and ecological traits a lesser extent, biological control using (Vargas 1970; Larrain 1986; Colomo et al. Bacillus thuringiensis, the egg parasitoid 2002; Luna et al. 2010). As an idiobiont Trichogramma pretiosum, and the predators species, it halts host development after Nesidiocoris tenuis and Podisus nigrispinus attacking it by the injection of venom. Upon (Miranda et al. 1998; Torres et al. 2002; eclosion, D. phtorimaeae larvae grow Journal of Insect Science | www.insectscience.org 2 Journal of Insect Science: Vol. 12 | Article 153 Savino et al. protected inside the leaf mines and pupate in non-concurrent, destructive host feeding situ. It is a solitary parasitoid that prefers the behavior, as shown by D. phtorimaeae, third-instar larval host to oviposit. One other particularly if associated to a synovigenic positive trait worth mentioning is its apparent strategy, could have a significant impact on its specificity for T. absoluta. ability to act as a successful biocontrol agent. To continue evaluating D. phtorimaeae Another point of interest in parasitoid biology potential as a biological control agent, its is the response of the ovipositional rate as a reproductive behavior and its functional function of host density, i.e., the functional Downloaded from https://academic.oup.com/jinsectscience/article-abstract/12/1/153/892634 by guest on 27 August 2019 response need to be addressed. Luna et al. response. This trait has been considered as (2010) found evidence that D. phtorimaeae one of the most important aspects in the practices non-concurrent destructive host dynamics of predator-prey interactions, and feeding, that is, the adult female consumes three main types are described: independency host haemolymph and tissues without (type I), direct (type III), and inverse density ovipositing, causing the death of the host dependency (type II) of parasitoid attack to (Jervis and Kidd 1986). Among the host density (Holling 1959; Hassell 2000). mechanisms described for this behavior, Particularly for biocontrol programs, punctures with the ovipositor and mouthparts functional response studies of potential and the construction of a feeding tube are candidates deal with the estimation of two mentioned. Occurrence of host feeding in parameters, the attack rate (a’) and the wasp parasitoids is related to ovigeny strategy, handling time (Th), to make comparisons or which can be pro-ovigenic (all eggs are predictions about possible control success. In mature at female’s emergence) or synovigenic a type I functional response, a’ is a constant (the female develops most of its lifetime egg proportion of killed hosts, in a type II it complement after emergence) (Jervis and gradually declines, and in a type III it linearly Copland 1996; Quicke 1997). Jervis et al. increases at low host densities and decreases (2001) devised an “ovigeny index,” which at high host densities (Juliano 2001; measures the degree of early-life Fernandez Arhex and Corley 2003; Luna et al. concentration of egg production. An index of 2007). Th is defined as the time needed to 1 refers to strict pro-ovigeny, whereas an manipulate the host and lay the eggs. Many index of 0 indicates extreme synovigeny. For parasitoids kill significant numbers of hosts idiobiont parasitoids, these authors pointed by feeding, besides parasitizing on them. For out a relationship between longer life parasitoids that practice non-concurrent, expectancy and host-feeding behavior, and destructive host
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