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Fitness performance of immatures under superparasitism and superparasitism strategy in an infanticidal semi-solitary parasitoid (Hymenoptera: Dryinidae): Effects of size of ovipositing females

HERLIN WERI 三重大学

2022.09.13

概要

Parasitoids, insects whose larvae feed and develop on or in insect hosts, often substantially influence the host population dynamics (Beddington et al. 1978). Therefore, many parasitoids have been used for biological control of pests (DeBach and Rosen 1991; Hajek 2004). In addition, parasitoids have been used to test various hypotheses proposed to explain foraging and sex allocation strategies in behavioral ecology (Godfray 1994; Wajnberg et al. 2008) because they are easy to rear and handle in the laboratory (see for 3.1. in chapter 3 for more details).

　Superparasitism–ovipositing in or on hosts that are parasitized by the same species (van Dijken and Waage 1987)–is common in parasitoids (Salt 1961; van Alphen and Visser 1990; Godfray 1994). It had been thought for a long time that superparasitism is a mistake of the ovipositing parasitoid–it occurs due to inability of the ovipositing female to discriminate parasitized hosts from unparasitized hosts–and that it is maladaptive behavior. However, superparasitism has been considered to be profitable, and adaptive under some situations; i.e., the parasitoid can get fitness gains through performing superparasitism, and should perform superparasitism under some situations (see for review van Alphen and Visser 1990; Speirs et al. 1991; Godfray 1994).

　Superparasitism is divided into self- and conspecific superparasitism: the former refers to when the first and second offspring (originating from the first and second ovipositions, respectively) come from the same mother, and the latter is when the two offsprings come from different mothers. When the survival rate of the second offspring is higher than zero, conspecific superparasitism rewards the mother with a fitness gain (van Alphen and Visser 1990). Meanwhile, self-superparasitism is generally less profitable than conspecific superparasitism due to the siblings facing competition for limited resources (e.g., Yamada and Miyamoto 1998; Yamada and Watanabe 2002; Yamada and Ikawa 2005; Zhang et al. 2014); in particular, it is usually non- or negatively profitable for solitary parasitoids except in the cases where multiple parasitoid immatures guarantee a higher emergence probability of one adult, which are probably created by the greater suppression of the immune systems of the host when multiple individuals are present (Puttler and van den Bosch 1959; Luna et al. 2016; Rasekh et al. 2018) or by the presence of conspecifics (van Alphen and Visser 1990; Yamada and Sugaura 2003; Ito and Yamada 2005, 2016).

　Body size has great influences on many physiological and ecological characteristics related to foraging, e.g., metabolic rate (Brown et al. 2004), food returned per foraging trip (Kerr et al. 2019), foraging range (Greenleaf et al. 2007; Weise et al. 2010; Orben et al. 2015), available foraging period (Streinzer et al. 2016), and searching efficiency in a patch (Visser 1994), and consequently prey (food)-foraging strategies are expected to change depending on the body size (Cozzoli et al. 2018; Josens et al. 2018). Prey foraging strategy usually aim to maximum energies obtained per time (Stephens and Krebs 1986), while oviposition strategy for parasitoids aims to maximum life-time fitness performance (Wajnberg et al. 2008). Thus, longevity and fecundity have also great influence on the oviposition strategy, and they are also determined primarily by body size (e.g., Heinz 1991; Visser 1994; West et al. 1996). Moreover, the mother size may have effects of the fitness performance of their offspring (mainly determined by the survival rate). Therefore, it must also influence decision-making related to oviposition. Unfortunately, the effects of the body size on the oviposition strategy and the fitness performance of the offspring have never been investigated so far in parasitoids, to my best knowledge.

　Whether superparasitism is adaptive is dependent on the fitness performance of offspring, physical conditions of the parasitoids, such as egg load and expected longevity, and the environmental situations in which the parasitoid is placed, such as host availability (van Alphen and Visser 1990; Speirs et al. 1991; Godfray 1994). Determination of the fitness performance of the first and second offspring under superparasitism is a basic step for understanding superparasitism strategy, including decision-making in superparasitism acceptance, sex allocation, selection of the oviposition place, and infanticide (ovicide and larvicide) for infanticidal parasitoids. However, few studies have revealed the fitness performance of the first and second offspring each under superparasitism (see for details 4.1. Introduction); in particular, the effects of the mother size of it have never been revealed.

　Here, I determined superparasitism strategy of the infanticidal parasitoid Echthrodelphax fairchildii Perkin (Hymenoptera: Dryinidae), using the fourth instar nymph of a host species, Laodelphax striatellus (Fallén) (Homoptera: Delphacidae); in particular, I focused the body-size effects on it. First, I determined the body-size effects of the first and second ovipositing females on the fitness performance of the first and second offspring separately under self and conspecific superparasitism with different intervals of the first and second ovipositions (called oviposition intervals hereafter) for the cases of occurrence and non-occurrence of infanticide. I also explored the possibility that the fitness performance of offspring is influenced by the mother size under single parasitism to understand the above body-size effect under superparasitism. Through these studies, I discovered the strong effects of the body size of the ovipositing females under single parasitism and superparasitism, as seen below. Then, I determined the frequencies of superparasitism acceptance and infanticide, selection of the oviposition place, and sex allocation under self and conspecific superparasitism with different oviposition intervals when the female parasitoid was placed in an environment with low availability of suitable hosts. In particular, I focused on the effects of the parasitoid size of the above decision-making.

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