482
tuffaceous shales (Ren et al. 2010b). Neuroptera consists of seven genera and nine
species of Mantispidae. Potentially parasitoid Coleoptera consisted of 2 families,
Staphylinidae and Carabidae, accounting for 21 genera and 37 species (Yu et al.
2019). The overwhelming majority of the beetles were Staphylinidae, although subfamily placement is tricky (Tan et al. 2010). Parasitoid Diptera include
Nemestrinimorpha (Nemestrinidae) and Eremochaetidae, consisting of two families, six genera and ten species (Han et al. 2019), generally consistent with the values from the earlier Karatau and Yanliao biotas. For Hymenoptera, the several,
major, symphytan lineages detailed in the previous three biotas are repeated in the
Jehol Biota. Hymenopteran parasitoid lineages are Ephialtitoidea (
†
Ephialtitidae),
Evanioidea (Aulacidae,
†
Baissidae, Evaniidae,
†
Praeaulacidae), Proctotrupoidea
(
†
Mesoserphidae, Heloridae, Pelecinidae, Roproniidae, Serphidae), Ichneumonoidea
(Ichneumonidae), and the aculeate Chrysidoidea (
†
“Bethylonymidae”) and
Vespoidea (Scoliidae) (Table 11.1). The increase to 13 hymenopteran families of the
Jehol Biota from the 8 or 9 occurrences from the previous three biotas is significant.
In addition, the presence of approximately 60 Jehol species assigned to the 13 families triples the species from the Purbeck Biota, which consist of about 20 species
allocated to 9 families. The Jehol data indicate that there was a quantitative increase
in the numbers of parasitoid families and species from Phase 1 to Phase 2 of
the MMPR.
11.6.7 Insect Faunas After the Mid-Mesozoic
Parasitoid Revolution
After phases 1 and 2 established the MMPR, Phase 3 increased the upward trend by
adding a considerable number of families of two lineages: proctotrupomorph
Hymenoptera and aculeate Hymenoptera (Fig. 11.19). During Phase 3, the trend
lines for all other lineages remained at the same relative levels as they did for Phase
2. In Phase 4, there were increases of families in proctotrupomorph Hymenoptera,
eremoneuran Diptera, Coleoptera and the earliest occurrence of valid Strepsiptera;
the levels of other lineages essentially held flat. These differences in representation
of the seven parasitoid lineages in phases 3 and 4 can be gleaned from four diverse
insect faunas that strategically sample every few tens of millions of years the Late
Cretaceous and Paleogene.
Some of these trends may be conditioned by taphonomic style, in which the history of the parasitoid fossil record is very asymmetric. Because of the relative hard
boundary of no biologically significant amber deposits occurring before approximately 135 million years ago (Labandeira 2014a), deposits before, during and after
the MMPR have a differing taphonomic cast, from which the primary data originates. Deposits prior to the MMPR—Madygen (237–220 Ma), Molteno
(237–228 Ma), Beishan–Shangtu (209–201 Ma) and Cow Branch biotas
(209–199 Ma)—consist only of compression-impression fossils. Deposits during
the MMPR―the Yanliao (165 Ma), Karatau (approximately 158–156 Ma), Purbeck
C. C. Labandeira and L. Li
tuffaceous shales (Ren et al. 2010b). Neuroptera consists of seven genera and nine
species of Mantispidae. Potentially parasitoid Coleoptera consisted of 2 families,
Staphylinidae and Carabidae, accounting for 21 genera and 37 species (Yu et al.
2019). The overwhelming majority of the beetles were Staphylinidae, although subfamily placement is tricky (Tan et al. 2010). Parasitoid Diptera include
Nemestrinimorpha (Nemestrinidae) and Eremochaetidae, consisting of two families, six genera and ten species (Han et al. 2019), generally consistent with the values from the earlier Karatau and Yanliao biotas. For Hymenoptera, the several,
major, symphytan lineages detailed in the previous three biotas are repeated in the
Jehol Biota. Hymenopteran parasitoid lineages are Ephialtitoidea (
†
Ephialtitidae),
Evanioidea (Aulacidae,
†
Baissidae, Evaniidae,
†
Praeaulacidae), Proctotrupoidea
(
†
Mesoserphidae, Heloridae, Pelecinidae, Roproniidae, Serphidae), Ichneumonoidea
(Ichneumonidae), and the aculeate Chrysidoidea (
†
“Bethylonymidae”) and
Vespoidea (Scoliidae) (Table 11.1). The increase to 13 hymenopteran families of the
Jehol Biota from the 8 or 9 occurrences from the previous three biotas is significant.
In addition, the presence of approximately 60 Jehol species assigned to the 13 families triples the species from the Purbeck Biota, which consist of about 20 species
allocated to 9 families. The Jehol data indicate that there was a quantitative increase
in the numbers of parasitoid families and species from Phase 1 to Phase 2 of
the MMPR.
11.6.7 Insect Faunas After the Mid-Mesozoic
Parasitoid Revolution
After phases 1 and 2 established the MMPR, Phase 3 increased the upward trend by
adding a considerable number of families of two lineages: proctotrupomorph
Hymenoptera and aculeate Hymenoptera (Fig. 11.19). During Phase 3, the trend
lines for all other lineages remained at the same relative levels as they did for Phase
2. In Phase 4, there were increases of families in proctotrupomorph Hymenoptera,
eremoneuran Diptera, Coleoptera and the earliest occurrence of valid Strepsiptera;
the levels of other lineages essentially held flat. These differences in representation
of the seven parasitoid lineages in phases 3 and 4 can be gleaned from four diverse
insect faunas that strategically sample every few tens of millions of years the Late
Cretaceous and Paleogene.
Some of these trends may be conditioned by taphonomic style, in which the history of the parasitoid fossil record is very asymmetric. Because of the relative hard
boundary of no biologically significant amber deposits occurring before approximately 135 million years ago (Labandeira 2014a), deposits before, during and after
the MMPR have a differing taphonomic cast, from which the primary data originates. Deposits prior to the MMPR—Madygen (237–220 Ma), Molteno
(237–228 Ma), Beishan–Shangtu (209–201 Ma) and Cow Branch biotas
(209–199 Ma)—consist only of compression-impression fossils. Deposits during
the MMPR―the Yanliao (165 Ma), Karatau (approximately 158–156 Ma), Purbeck
C. C. Labandeira and L. Li