479
fine-grained shales (Grimaldi and Engel 2005a). The Solite Quarry has not yielded
Hymenoptera. However, Diptera are diverse compared to other Late Triassic to earliest Jurassic intervals, consisting of the nematocerous Tipulomorpha (
†
Vladipteridae,
Limoniidae), Psychodomorpha (Psychodidae,
†
Eoptychopteridae, an undetermined
family),
Culicomorpha
(
†
undetermined
family),
Bibionomorpha
(
†
Procramptonomyiidae,
†
Protorhyphidae,
†
Paraxymyiidae,
†
Crossaphididae), and
stem-group Brachycera (
†
Prosechamyiidae; Blagoderov et al. 2007). Although the
undetermined family of Culicomorpha could have been a parasite as a blood feeder
(Fig. 11.8; Blagoderov et al. 2007), all other evidence from subsequent fossil lineages and modern biology would disallow a parasitoid interpretation for this culicomorph. The Solite Coleoptera includes Staphylinidae (Fraser et al. 1996), a rove
beetle, which almost certainly did not belong to a subclade such as the Aleocharinae
that includes modern parasitoid members (Klimaszewski 1984).
11.6.6 Insect Faunas During the Mid-Mesozoic
Parasitoid Revolution
Four of the most informative insect faunas for documenting the initial expansion
interval of the MMPR during phases 1 and 2 range from the latest Middle Jurassic
(late Callovian Stage) to the mid Early Cretaceous (mid Albian Stage), before the
shift from phase 2 to phase 3. The deposits, from oldest to youngest, originate from
northeastern China, south-central Kazakhstan, southern England in the United
Kingdom, and back to northeastern China. Insect taxa from these four deposits collectively exhibit (1) the first appearances and diversification and ecological expansion of non-proctotrupomorph and proctotrupomorph hymenopteran parasitoids;
(2) to a lesser extent, the expansion of non-Eremoneura Diptera and Coleoptera; and
(3) the modest beginnings of the Aculeata Hymenoptera and perhaps Strepsiptera
diversifications. The oldest Early Cretaceous amber deposits such as Lebanese and
Álava amber are undoubtedly extensions of Phase 2 of the MMPR, but their parasitoid faunas remain largely unstudied. By the end of phases 1 and 2 of the parasitoid
diversification events, six of the seven major clades of parasitoids were established;
the exception was Strepsiptera.
The first two insect faunas are Jurassic in age and they provide documentation
for diversification of the earliest lineages of parasitoids. The first fauna is from the
Jiulongshan Formation, representing the Yanliao Biota, and located in the Daohugou
area where the three provinces of Liaoning, Hebei and Inner Mongolia come
together (Ren et al. 2010a; Huang 2016). The age of the Yanliao Biota is 165 Ma,
based on secure radioisotopic age dates (Ren et al. 2010b; Huang et al. 2016),
and equivalent to the late part of the Callovian Stage (Walker et al. 2013). Yanliao
fossils occur mostly in medium to dark hued tuffaceous siltstones that were deposited in fluvial and geographically extensive lacustrine environments (Ren et al.
2010b). By the time the Jiulongshan Formation was deposited, there is evidence for
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
fine-grained shales (Grimaldi and Engel 2005a). The Solite Quarry has not yielded
Hymenoptera. However, Diptera are diverse compared to other Late Triassic to earliest Jurassic intervals, consisting of the nematocerous Tipulomorpha (
†
Vladipteridae,
Limoniidae), Psychodomorpha (Psychodidae,
†
Eoptychopteridae, an undetermined
family),
Culicomorpha
(
†
undetermined
family),
Bibionomorpha
(
†
Procramptonomyiidae,
†
Protorhyphidae,
†
Paraxymyiidae,
†
Crossaphididae), and
stem-group Brachycera (
†
Prosechamyiidae; Blagoderov et al. 2007). Although the
undetermined family of Culicomorpha could have been a parasite as a blood feeder
(Fig. 11.8; Blagoderov et al. 2007), all other evidence from subsequent fossil lineages and modern biology would disallow a parasitoid interpretation for this culicomorph. The Solite Coleoptera includes Staphylinidae (Fraser et al. 1996), a rove
beetle, which almost certainly did not belong to a subclade such as the Aleocharinae
that includes modern parasitoid members (Klimaszewski 1984).
11.6.6 Insect Faunas During the Mid-Mesozoic
Parasitoid Revolution
Four of the most informative insect faunas for documenting the initial expansion
interval of the MMPR during phases 1 and 2 range from the latest Middle Jurassic
(late Callovian Stage) to the mid Early Cretaceous (mid Albian Stage), before the
shift from phase 2 to phase 3. The deposits, from oldest to youngest, originate from
northeastern China, south-central Kazakhstan, southern England in the United
Kingdom, and back to northeastern China. Insect taxa from these four deposits collectively exhibit (1) the first appearances and diversification and ecological expansion of non-proctotrupomorph and proctotrupomorph hymenopteran parasitoids;
(2) to a lesser extent, the expansion of non-Eremoneura Diptera and Coleoptera; and
(3) the modest beginnings of the Aculeata Hymenoptera and perhaps Strepsiptera
diversifications. The oldest Early Cretaceous amber deposits such as Lebanese and
Álava amber are undoubtedly extensions of Phase 2 of the MMPR, but their parasitoid faunas remain largely unstudied. By the end of phases 1 and 2 of the parasitoid
diversification events, six of the seven major clades of parasitoids were established;
the exception was Strepsiptera.
The first two insect faunas are Jurassic in age and they provide documentation
for diversification of the earliest lineages of parasitoids. The first fauna is from the
Jiulongshan Formation, representing the Yanliao Biota, and located in the Daohugou
area where the three provinces of Liaoning, Hebei and Inner Mongolia come
together (Ren et al. 2010a; Huang 2016). The age of the Yanliao Biota is 165 Ma,
based on secure radioisotopic age dates (Ren et al. 2010b; Huang et al. 2016),
and equivalent to the late part of the Callovian Stage (Walker et al. 2013). Yanliao
fossils occur mostly in medium to dark hued tuffaceous siltstones that were deposited in fluvial and geographically extensive lacustrine environments (Ren et al.
2010b). By the time the Jiulongshan Formation was deposited, there is evidence for
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution