385
hyperparasitoids exhibit very high host specificity (Schär and Vorburger 2013). The
concept of hyperparasitoidism is inextricably linked to hyperparasitism in the older
biological literature, where the two life histories were not distinguished. The French
naturalist Maria Sibylla Merian (1647–1717) was probably the first person to recognize hyperparasitoidism (as hyperparasitism), as illustrated in her drawings (Todd
2011). The concept later was recognized in verse in 1733 by the English writer
Jonathan Swift, who undoubtedly borrowed the idea from earlier authors.
11.2.5.6 Egg Parasitoidism, Larval Parasitoidism, Pupal Parasitoidism
and Adult Parasitoidism
Parasitoids of holometabolous insects can attack any of the four major developmental stages, or any two of adjacent stages of the egg, multiple instars of the larva, pupa
and adult. Frequently parasitoids are not only species specific but also target particular developmental instars, such as egg parasitoids (Malyshev 1968; Huber 1986;
Whitfield 1998), larval parasitoids (Askew 1971; Weinstein and Austin 1991;
Whitfield 2003), pupal parasitoids (Clausen 1940; Quevillon and Hughes 2018) and
adult parasitoids (Askew 1971; Whitfield 1998, 2003), although it is the earlier
stages that are most often attacked. For hemimetabolous insect hosts, parasitoids
attack their eggs (Muldrew 1953; Greathead 1963; Brown 1973); nymphs, if terrestrial (Eggleton and Belshaw 1992; Cook 2014); naiads, if aquatic (Clausen 1940;
Askew 1971); or adults (DeBach 1964; Kirkpatrick 1947; Elzinga 1977; Cook
2014), although there is little somatic differentiation between nymphal and their
conspecific adult instars. Parasitoids of egg, pupal and adult stages of their hosts
typically are idiobionts, as are those parasitoids of larvae whose sting causes permanent paralysis (Gauld and Bolton 1988; Godfray 1994). Koinobiont parasitoids are
those that attack the egg–larval and larval–pupal instar couplets or are parasitoids of
adjacent larval instars, and do not paralyze their hosts. A special category is an egg
parasitoid, which often is a miniscule insect that deposits typically very small eggs
on the surface or in the interstitial tissues of a much larger host egg. Other egg parasitoids oviposit in an egg sac containing multiple eggs, in which the hatched parasitoid larva either slowly consumes nutritive tissues of the egg that eventually
starves and kills the embryo, or alternatively successively consumes eggs within an
egg sac (Askew 1971; Vetter et al. 2012).
11.2.5.7 Cleptoparasitoidism
Cleptoparasitoidism is a rare life style in which a parasitoid absconds a vital resource
such as food that results in the death of the host after it is deprived of the supply
(Frank and Gillett-Kaufman 2006; Dehon et al. 2017). Food resources can include
foliage, galls, insects or other provisions intended for the host (Eggleton and
Belshaw 1992). A related concept is social parasitism, which is a special type of
parasitism constituting a relationship between two species such that one species
is dependent parasitically on the other (Brandt et al. 2005; Smith et al. 2007).
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
hyperparasitoids exhibit very high host specificity (Schär and Vorburger 2013). The
concept of hyperparasitoidism is inextricably linked to hyperparasitism in the older
biological literature, where the two life histories were not distinguished. The French
naturalist Maria Sibylla Merian (1647–1717) was probably the first person to recognize hyperparasitoidism (as hyperparasitism), as illustrated in her drawings (Todd
2011). The concept later was recognized in verse in 1733 by the English writer
Jonathan Swift, who undoubtedly borrowed the idea from earlier authors.
11.2.5.6 Egg Parasitoidism, Larval Parasitoidism, Pupal Parasitoidism
and Adult Parasitoidism
Parasitoids of holometabolous insects can attack any of the four major developmental stages, or any two of adjacent stages of the egg, multiple instars of the larva, pupa
and adult. Frequently parasitoids are not only species specific but also target particular developmental instars, such as egg parasitoids (Malyshev 1968; Huber 1986;
Whitfield 1998), larval parasitoids (Askew 1971; Weinstein and Austin 1991;
Whitfield 2003), pupal parasitoids (Clausen 1940; Quevillon and Hughes 2018) and
adult parasitoids (Askew 1971; Whitfield 1998, 2003), although it is the earlier
stages that are most often attacked. For hemimetabolous insect hosts, parasitoids
attack their eggs (Muldrew 1953; Greathead 1963; Brown 1973); nymphs, if terrestrial (Eggleton and Belshaw 1992; Cook 2014); naiads, if aquatic (Clausen 1940;
Askew 1971); or adults (DeBach 1964; Kirkpatrick 1947; Elzinga 1977; Cook
2014), although there is little somatic differentiation between nymphal and their
conspecific adult instars. Parasitoids of egg, pupal and adult stages of their hosts
typically are idiobionts, as are those parasitoids of larvae whose sting causes permanent paralysis (Gauld and Bolton 1988; Godfray 1994). Koinobiont parasitoids are
those that attack the egg–larval and larval–pupal instar couplets or are parasitoids of
adjacent larval instars, and do not paralyze their hosts. A special category is an egg
parasitoid, which often is a miniscule insect that deposits typically very small eggs
on the surface or in the interstitial tissues of a much larger host egg. Other egg parasitoids oviposit in an egg sac containing multiple eggs, in which the hatched parasitoid larva either slowly consumes nutritive tissues of the egg that eventually
starves and kills the embryo, or alternatively successively consumes eggs within an
egg sac (Askew 1971; Vetter et al. 2012).
11.2.5.7 Cleptoparasitoidism
Cleptoparasitoidism is a rare life style in which a parasitoid absconds a vital resource
such as food that results in the death of the host after it is deprived of the supply
(Frank and Gillett-Kaufman 2006; Dehon et al. 2017). Food resources can include
foliage, galls, insects or other provisions intended for the host (Eggleton and
Belshaw 1992). A related concept is social parasitism, which is a special type of
parasitism constituting a relationship between two species such that one species
is dependent parasitically on the other (Brandt et al. 2005; Smith et al. 2007).
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution