453
contrast, the parasitoid habit evolved into predation on bees in soil and plant litter
habitats and possibly on grasshopper egg pod predation. The oldest fossils of
Cleridae are from the latest Middle Jurassic of China (Kolibáč and Huang 2016).
Bothrideridae (dry bark beetles) consists of 38 genera, 400 species and 4 subfamilies, where one subfamily, Bothriderinae, has ectoparasitoid life habits (Philips
and Ivie 2002). Bothrideridae are highly elongate beetles whose adults occur under
tree bark or in composted soil. Their nonparasitic larvae feed on fungi whereas their
hypermetamorphic, host-seeking, first-instar larvae is a spinose triungulin that
attacks and becomes an ectoparasitoid on wood-boring larvae of metallic woodboring beetles, ambrosia beetles, bark beetles, deathwatch beetles, augur beetles,
longhorn beetles, and wood wasps (Horion 1961; Ślipiński et al. 2010). Subsequent
larval instars are grub-like and complete an ectoparasitoid existence on their host
(Philips and Ivie 2002; Lawrence and Ślipiński 2013). The ectoparasite habit likely
originated from mycophagy in an ancestral bothriderid lineage (Crowson 1981),
and subsequently evolved into ectoparasitoidism. Ectoparasitoidism currently is
confined to the diverse Subfamily Bothriderinae, where adults of several genera
bear lightly sclerotized, swollen abdomens probably involved in very fecund levels
of egg production. The oldest Bothrideridae are from the middle Eocene
(Ponomarenko 1995).
Passandridae (flat bark beetles) consist of 9 genera and 109 species that are ectoparasitoids of the larvae or pupae of wood boring insects such as weevils, longhorned beetles, ambrosia beetles, bark beetles or wood wasps that live in dead wood
or under bark (Crowson 1981; Thomas 2002; Burkhardt and Ślipiński 2010).
Passandrids also assume a second major parasitoid strategy as hyperparasitoids on
braconid wasps (Burkhardt and Ślipiński 2010). The larval body is highly modified
for ectoparasitoid habits and display changes in larval morphology from instar to
instar (Burkhardt and Ślipiński 2003), similar to hypermetamorphic larvae.
Passandrid larval morphology externally is variously sclerotized, flattened and
spiny, with an enlarged abdomen and reduced mouthparts. Evidently, parasitoid larvae of Passandridae display limited host specificity. Adults are considerably flat
dorsoventrally and occupy subcortical bark habitats. The oldest fossils of
Passandridae evidently are Cenomanian in age (Ponomarenko 1995).
Meloidae (blister beetles) are mostly parasitoids of grasshoppers and nondomesticated bees (Lawrence et al. 1991; Arnett Jr et al. 2002). Meloid first-instar
triungulin larvae frequently are encountered on flowers, are acquired by foraging
male bees, and then are transported to female bees during mating (Lawrence et al.
1991; Arnett Jr et al. 2002). At the bee nest site, the triungulin larvae invade bee
cells where eggs reside and successively consume either the egg or the developing
larva. Meloid larvae undergo hypermetamorphosis and initially are ambulatory,
host-seeking triungulins, followed by sessile, grub-like larval instars that emerge
from their host insect nest as adult blister beetles.
About 120 genera and 2500 species of Meloidae have larvae that are predaceous
or more frequently parasitoids (Bologna et al. 2010). Larval hypermetamorphosis is
common and involves host-searching behavior, and after attack of the host, the larva
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
contrast, the parasitoid habit evolved into predation on bees in soil and plant litter
habitats and possibly on grasshopper egg pod predation. The oldest fossils of
Cleridae are from the latest Middle Jurassic of China (Kolibáč and Huang 2016).
Bothrideridae (dry bark beetles) consists of 38 genera, 400 species and 4 subfamilies, where one subfamily, Bothriderinae, has ectoparasitoid life habits (Philips
and Ivie 2002). Bothrideridae are highly elongate beetles whose adults occur under
tree bark or in composted soil. Their nonparasitic larvae feed on fungi whereas their
hypermetamorphic, host-seeking, first-instar larvae is a spinose triungulin that
attacks and becomes an ectoparasitoid on wood-boring larvae of metallic woodboring beetles, ambrosia beetles, bark beetles, deathwatch beetles, augur beetles,
longhorn beetles, and wood wasps (Horion 1961; Ślipiński et al. 2010). Subsequent
larval instars are grub-like and complete an ectoparasitoid existence on their host
(Philips and Ivie 2002; Lawrence and Ślipiński 2013). The ectoparasite habit likely
originated from mycophagy in an ancestral bothriderid lineage (Crowson 1981),
and subsequently evolved into ectoparasitoidism. Ectoparasitoidism currently is
confined to the diverse Subfamily Bothriderinae, where adults of several genera
bear lightly sclerotized, swollen abdomens probably involved in very fecund levels
of egg production. The oldest Bothrideridae are from the middle Eocene
(Ponomarenko 1995).
Passandridae (flat bark beetles) consist of 9 genera and 109 species that are ectoparasitoids of the larvae or pupae of wood boring insects such as weevils, longhorned beetles, ambrosia beetles, bark beetles or wood wasps that live in dead wood
or under bark (Crowson 1981; Thomas 2002; Burkhardt and Ślipiński 2010).
Passandrids also assume a second major parasitoid strategy as hyperparasitoids on
braconid wasps (Burkhardt and Ślipiński 2010). The larval body is highly modified
for ectoparasitoid habits and display changes in larval morphology from instar to
instar (Burkhardt and Ślipiński 2003), similar to hypermetamorphic larvae.
Passandrid larval morphology externally is variously sclerotized, flattened and
spiny, with an enlarged abdomen and reduced mouthparts. Evidently, parasitoid larvae of Passandridae display limited host specificity. Adults are considerably flat
dorsoventrally and occupy subcortical bark habitats. The oldest fossils of
Passandridae evidently are Cenomanian in age (Ponomarenko 1995).
Meloidae (blister beetles) are mostly parasitoids of grasshoppers and nondomesticated bees (Lawrence et al. 1991; Arnett Jr et al. 2002). Meloid first-instar
triungulin larvae frequently are encountered on flowers, are acquired by foraging
male bees, and then are transported to female bees during mating (Lawrence et al.
1991; Arnett Jr et al. 2002). At the bee nest site, the triungulin larvae invade bee
cells where eggs reside and successively consume either the egg or the developing
larva. Meloid larvae undergo hypermetamorphosis and initially are ambulatory,
host-seeking triungulins, followed by sessile, grub-like larval instars that emerge
from their host insect nest as adult blister beetles.
About 120 genera and 2500 species of Meloidae have larvae that are predaceous
or more frequently parasitoids (Bologna et al. 2010). Larval hypermetamorphosis is
common and involves host-searching behavior, and after attack of the host, the larva
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution