480
the commencement of a major evolutionary expansion of hymenopteran parasitoid
lineages. Probable parasitoids of Neuroptera consisted of one family, Mantispidae,
with two genera and two species. Possible parasitoid taxa of Coleoptera may have
been present, representing two families, Staphylinidae and Ripiphoridae, four genera and four species (Tan et al. 2010), indicating that beetles were a very minor
component of the parasitoid guild. For Staphylinidae, there is no evidence that any
subfamilies known to house parasitoid taxa, such as Aleocharinae, were present
during the late Middle Jurassic. Dipteran parasitoid taxa were less diverse, consisting of Nemestrinimorpha (Nemestrinidae) and Eremochaetidae representing two
families, five genera and eight species (Zhang et al. 2017; Ren et al. 2019), marking
the early presence of dipteran parasitoids in the MMPR. Well-represented
hymenopteran taxa include a diversity of plant-associated sawflies (“symphytans”)
that are placed in the superfamilies Xyeloidea (Xyelidae,
†
Daohugoidae),
Tenthredinoidea (
†
Xyelotomidae), Pamphilioidea (
†
Xyelydidae), Cephoidea
(
†
Sepulcidae), Siricoidea (Anaxyelidae, Siricidae), and the parasitoid Orussoidea
(
†
Karatavitidae) that shares a sister-group relationship with parasitoid Apocrita.
Apocritan parasitoids occur in the three superfamilies Evanioidea (
†
Anomopterellidae,
†
Praeaulacidae), Ephialtitoidea (
†
Ephialtitidae) and Proctotrupoidea (Heloridae,
†
Mesoserphidae, Pelecinidae, Roproniidae) that collectively account for approximately 15 genera (Rasnitsyn and Zhang 2004, 2010; Gao et al. 2010; Huang and Cai
2016; Li et al. 2018a; Wang et al. 2019; Table 11.1). This distribution of hymenopteran
parasitoid families indicates that eight major, family-level lineages of orussoid sawflies and apocritan parasitoid wasps were present, extending into the basal
Proctotrupomorpha superclade during Yanliao time (Fig. 11.19). The Jiulongshan
Formation probably represents the earliest, most extensive sample of Phase 1 of the
MMPR of any biota worldwide.
The second major insect fauna is the Karabastau Formation, representing the
Karatau Biota, located along the Karatau Range, near the towns of Aulie (formerly
Mikhailovka) and Uspenovka (formerly Galkino) in southern Kazakhstan, and considered Late Jurassic in age. Because of the multiple localities representing multiple
stratigraphic horizons at Karatau, the deposits are assigned to a time range that
includes the Oxfordian and Kimmeridgian stages (Rasnitsyn and Zherikhin 2002),
corresponding to 164–152 Ma (Walker et al. 2013), making the Karatau Biota seven
million years younger, on average, than the Yanliao Biota. The insect fossils occur
in dark grey shales that preserve detail such as wing eyespots, delicate surface ornamentation and fine hairs on body surfaces (Rohdendorf 1968a; Grimaldi and Engel
2005a). Neuroptera consists of only one genus and species of Mantispidae.
Concerning the parasitoid taxa, Coleoptera consisted of 2 families, Staphylinidae
and Anthribidae, consisting of 10 genera and 12 species, although Staphylinidae at
Karatau have not been assigned to the extant parasitoid subfamily Aleocharinae
(Tichomirova 1968; Yu et al. 2019), which contains many modern parasitoid species
(Klimaszewski 1984). Parasitoid Diptera include Nemestrinimorpha (Nemestrinidae
and Acroceridae) and Archisargoidea (Eremochaetidae), consisting of 3 families, 8
genera and 16 species of potentially parasitoidic flies (Rohdendorf 1968b; Mostovski
1998; Zhang et al. 2016) (Table 11.1) comparable in diversity to that of the Yanliao
C. C. Labandeira and L. Li
the commencement of a major evolutionary expansion of hymenopteran parasitoid
lineages. Probable parasitoids of Neuroptera consisted of one family, Mantispidae,
with two genera and two species. Possible parasitoid taxa of Coleoptera may have
been present, representing two families, Staphylinidae and Ripiphoridae, four genera and four species (Tan et al. 2010), indicating that beetles were a very minor
component of the parasitoid guild. For Staphylinidae, there is no evidence that any
subfamilies known to house parasitoid taxa, such as Aleocharinae, were present
during the late Middle Jurassic. Dipteran parasitoid taxa were less diverse, consisting of Nemestrinimorpha (Nemestrinidae) and Eremochaetidae representing two
families, five genera and eight species (Zhang et al. 2017; Ren et al. 2019), marking
the early presence of dipteran parasitoids in the MMPR. Well-represented
hymenopteran taxa include a diversity of plant-associated sawflies (“symphytans”)
that are placed in the superfamilies Xyeloidea (Xyelidae,
†
Daohugoidae),
Tenthredinoidea (
†
Xyelotomidae), Pamphilioidea (
†
Xyelydidae), Cephoidea
(
†
Sepulcidae), Siricoidea (Anaxyelidae, Siricidae), and the parasitoid Orussoidea
(
†
Karatavitidae) that shares a sister-group relationship with parasitoid Apocrita.
Apocritan parasitoids occur in the three superfamilies Evanioidea (
†
Anomopterellidae,
†
Praeaulacidae), Ephialtitoidea (
†
Ephialtitidae) and Proctotrupoidea (Heloridae,
†
Mesoserphidae, Pelecinidae, Roproniidae) that collectively account for approximately 15 genera (Rasnitsyn and Zhang 2004, 2010; Gao et al. 2010; Huang and Cai
2016; Li et al. 2018a; Wang et al. 2019; Table 11.1). This distribution of hymenopteran
parasitoid families indicates that eight major, family-level lineages of orussoid sawflies and apocritan parasitoid wasps were present, extending into the basal
Proctotrupomorpha superclade during Yanliao time (Fig. 11.19). The Jiulongshan
Formation probably represents the earliest, most extensive sample of Phase 1 of the
MMPR of any biota worldwide.
The second major insect fauna is the Karabastau Formation, representing the
Karatau Biota, located along the Karatau Range, near the towns of Aulie (formerly
Mikhailovka) and Uspenovka (formerly Galkino) in southern Kazakhstan, and considered Late Jurassic in age. Because of the multiple localities representing multiple
stratigraphic horizons at Karatau, the deposits are assigned to a time range that
includes the Oxfordian and Kimmeridgian stages (Rasnitsyn and Zherikhin 2002),
corresponding to 164–152 Ma (Walker et al. 2013), making the Karatau Biota seven
million years younger, on average, than the Yanliao Biota. The insect fossils occur
in dark grey shales that preserve detail such as wing eyespots, delicate surface ornamentation and fine hairs on body surfaces (Rohdendorf 1968a; Grimaldi and Engel
2005a). Neuroptera consists of only one genus and species of Mantispidae.
Concerning the parasitoid taxa, Coleoptera consisted of 2 families, Staphylinidae
and Anthribidae, consisting of 10 genera and 12 species, although Staphylinidae at
Karatau have not been assigned to the extant parasitoid subfamily Aleocharinae
(Tichomirova 1968; Yu et al. 2019), which contains many modern parasitoid species
(Klimaszewski 1984). Parasitoid Diptera include Nemestrinimorpha (Nemestrinidae
and Acroceridae) and Archisargoidea (Eremochaetidae), consisting of 3 families, 8
genera and 16 species of potentially parasitoidic flies (Rohdendorf 1968b; Mostovski
1998; Zhang et al. 2016) (Table 11.1) comparable in diversity to that of the Yanliao
C. C. Labandeira and L. Li