463
is known only for one specimen attached to a leafhopper (Auchenorrhyncha:
Cicadellidae) from the early Miocene of the Dominican Republic (Poinar Jr and
Poinar 1999). Analogously, Pyralidae (snout moths) contain a single species,
Sthenauge parasitus, which is an ectoparasitoid on the larvae of the saturniid moth
Aplomerus (Lepidoptera: Saturniidae), initially feeding on the host’s dermal spines
but eventually consuming it. Known parasitoidism arose at least twice in Lepidoptera,
separately in Epipyropidae and Pyralidae, but likely arose many more times, particularly within lineages of Epipyropidae, given the poor understanding of the lifehabits and phylogeny of these moths (Jeon et al. 2002). Parasitoidism in Epipyropidae
plausibly evolved from ectoparasitic behavior occurring on the hemipteran host surface, followed by entering the host by piercing its cuticle and then eventually killing
it with the exit of the parasite larva (Eggleton and Belshaw 1992). The evolutionary
source of parasitoidism in Epipyropidae and Pyralidae is unclear, and there is no
fossil record for parasitoid members of either lineage.
11.5.2.7 Hymenoptera (Wasps)
Hymenoptera constitute about 75% of all modern parasitoid species and have the
most diverse taxonomic spectrum of parasitoid lineages of any order of insects
(Santos and Quicke 2011). The 17 hymenopteran superfamilies of parasitoids are
approximately split between ectoparasitoid and endoparasitoid dominated superfamilies; they include 4 superfamilies that contain hyperparasitoids, 4 superfamilies
that have predators, and 2 superfamilies with cleptoparasitoids (Gauld and Bolton
1988) (Table 11.1). One superfamily, Chalcidoidea, while overwhelmingly dominated by parasitoid taxa, contains two families, Agaonidae and Chalcididae, which
are dominantly represented by taxa such as pollinators, gallers, seed predators and
herbivores as well as cleptoparasitoids in plant galls (Eggleton and Belshaw 1992).
Family-level diversity per superfamily ranges from 1 to 20. There are 92 families of
extant and extinct hymenopteran parasitoids, representing 63% of all insect families
with parasitoid life habits (Table 11.2). Of these families, 22 (23.9%) are extinct,
some of which represent stem-group lineages. Even though there are 3.75 times
more hymenopteran than dipteran parasitoid species, hymenopteran hosts are
restricted to one phylum, Arthropoda, and within that phylum only Insecta are
attacked (Morris 1998). One possibility for this vast disparity in host utilization is
that, whereas parasitoidism arose once or twice in Hymenoptera (Dowton and
Austin 1995b). It probably arose at least 10 to 100 times in Diptera (Eggleton and
Belshaw 1992). Evidently, competition for the variety of targeted hosts was much
less constrained in Diptera.
Sawflies (Tenthredinoidea, Siricoidea) are the sister-group to the parasitoid
clades Orussidae and Apocrita, which represent the single (Gauld and Bolton 1988)
or dual (Dowton and Austin 1995b) origins of parasitoidism within Hymenoptera. It
appears that the parasitoid condition in Hymenoptera is associated with a founder
effect of considerably higher content of adenine and thymine, as opposed to cytosine and guanine, in the mitochondrial DNA of Apocrita wasps, when compared to
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
is known only for one specimen attached to a leafhopper (Auchenorrhyncha:
Cicadellidae) from the early Miocene of the Dominican Republic (Poinar Jr and
Poinar 1999). Analogously, Pyralidae (snout moths) contain a single species,
Sthenauge parasitus, which is an ectoparasitoid on the larvae of the saturniid moth
Aplomerus (Lepidoptera: Saturniidae), initially feeding on the host’s dermal spines
but eventually consuming it. Known parasitoidism arose at least twice in Lepidoptera,
separately in Epipyropidae and Pyralidae, but likely arose many more times, particularly within lineages of Epipyropidae, given the poor understanding of the lifehabits and phylogeny of these moths (Jeon et al. 2002). Parasitoidism in Epipyropidae
plausibly evolved from ectoparasitic behavior occurring on the hemipteran host surface, followed by entering the host by piercing its cuticle and then eventually killing
it with the exit of the parasite larva (Eggleton and Belshaw 1992). The evolutionary
source of parasitoidism in Epipyropidae and Pyralidae is unclear, and there is no
fossil record for parasitoid members of either lineage.
11.5.2.7 Hymenoptera (Wasps)
Hymenoptera constitute about 75% of all modern parasitoid species and have the
most diverse taxonomic spectrum of parasitoid lineages of any order of insects
(Santos and Quicke 2011). The 17 hymenopteran superfamilies of parasitoids are
approximately split between ectoparasitoid and endoparasitoid dominated superfamilies; they include 4 superfamilies that contain hyperparasitoids, 4 superfamilies
that have predators, and 2 superfamilies with cleptoparasitoids (Gauld and Bolton
1988) (Table 11.1). One superfamily, Chalcidoidea, while overwhelmingly dominated by parasitoid taxa, contains two families, Agaonidae and Chalcididae, which
are dominantly represented by taxa such as pollinators, gallers, seed predators and
herbivores as well as cleptoparasitoids in plant galls (Eggleton and Belshaw 1992).
Family-level diversity per superfamily ranges from 1 to 20. There are 92 families of
extant and extinct hymenopteran parasitoids, representing 63% of all insect families
with parasitoid life habits (Table 11.2). Of these families, 22 (23.9%) are extinct,
some of which represent stem-group lineages. Even though there are 3.75 times
more hymenopteran than dipteran parasitoid species, hymenopteran hosts are
restricted to one phylum, Arthropoda, and within that phylum only Insecta are
attacked (Morris 1998). One possibility for this vast disparity in host utilization is
that, whereas parasitoidism arose once or twice in Hymenoptera (Dowton and
Austin 1995b). It probably arose at least 10 to 100 times in Diptera (Eggleton and
Belshaw 1992). Evidently, competition for the variety of targeted hosts was much
less constrained in Diptera.
Sawflies (Tenthredinoidea, Siricoidea) are the sister-group to the parasitoid
clades Orussidae and Apocrita, which represent the single (Gauld and Bolton 1988)
or dual (Dowton and Austin 1995b) origins of parasitoidism within Hymenoptera. It
appears that the parasitoid condition in Hymenoptera is associated with a founder
effect of considerably higher content of adenine and thymine, as opposed to cytosine and guanine, in the mitochondrial DNA of Apocrita wasps, when compared to
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution