481
Biota. No evidence exists for parasitoid Strepsiptera, Trichoptera or Lepidoptera.
Hymenoptera included a similar spectrum of sawflies (Rasnitsyn 1968) and parasitoid lineages that were more diverse than those in the Yanliao Biota. The hymenopteran
parasitoid lineages include the parasitoid sawfly Orussoidea (
†
Karatavitidae), and
parasitoid wasps Ephialtitoidea (
†
Ephialtitidae), Megalyroidea (Megalyridae),
Evanioidea (
†
Anomopterellidae,
†
Praeaulacidae), Proctotrupoidea (Heloridae,
†
Mesoserphidae) and the basal aculeate lineage Chrysidoidea (
†
“Bethylonymidae”)
(Kozlov 1968; Rasnitsyn 2002) (Table 11.1). The same number of eight, familylevel lineages of hymenopteran parasitoids was present in the Karatau Biota as in
the Yanliao Biota, but with two family substitutions and a doubling of species to
approximately 30, indicating evolutionary stability at the family level but an
increased proliferation of species. In general, it appears that parasitoids of the
Karatau Biota represent continued stability of major parasitoid lineages when compared with the Yanliao Biota, but increased generation of genera and species at
lower taxonomic levels.
The third insect fauna considered during the initial expansion of the MMPR
comes from the Lulworth Formation, constituting the lower unit of the Purbeck
Limestone Group, and its upper unit, the Durlston Formation, which crop out along
the Vale of Wardour area in Dorset, southern England (Rasnitsyn et al. 1998; Coram
et al. 2000; Coram and Jepson 2012). The age of the Purbeck strata is earliest
Cretaceous (Berriasian Stage), with a corresponding age date of 145–139 Ma
(Walker et al. 2013), and is approximately 16 million years younger than the Karatau
Biota. The strata of the Purbeck Biota containing the insect fossils consist of finegrained, mostly thinly bedded limestone occasionally interrupted by algal mat layers (Coram 2003). Parasitoid-containing lineages of Neuroptera, Coleoptera,
Strepsiptera, Diptera, Trichoptera, and Lepidoptera are absent. However, as in the
preceding two fossil units, a near full complement of sawflies is present, excluding
the Orussoidea—the only parasitoid sawfly clade. Apocritan wasps consist of
Megalyroidea (Megalyridae), Evanioidea (cf. Aulacidae,
†
Baissidae,
†
incertae
sedis),
Proctotrupoidea
(Diapriidae,
Proctotrupidae),
Ichneumonoidea
(Ichneumonidae), the aculeate lineages Chrysidoidea (
†
“Bethylonymidae”) and
Apoidea (Sphecidae), and the undetermined apocritan Apocrites (Coram and Jepson
2012) (Table 11.1). Including unassigned but distinctive lineages, there are 10
family- level lineages of hymenopteran parasitoids representing about 20 species in
the Purbeck Biota.
The Yixian Formation is the fourth insect fauna that highlights Phase 2 of the
MMPR (Fig. 11.19). Fossils from the Yixian Formation formed the distinctive Jehol
Biota that originate from seven or eight major, fossil-yielding localities, mostly in
Liaoning Province of northeastern China, west of Beipiao City and near Liaodong
Bay, an arm of the Yellow Sea (Zhang et al. 2010). Although once controversial
(Ren et al. 2010b), the date currently is established as mid Early Cretaceous, from
126 to 122 Ma (Swisher III et al. 1999), equivalent to late Barremian to early Aptian
in age (Walker et al. 2013), and approximately 18 million years younger than the
Purbeck Biota. The fossils occur in broad outcrops of lake deposits with lightcolored, highly oxidized, fine-grained mudstone and siltstone strata of buff-colored
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
Biota. No evidence exists for parasitoid Strepsiptera, Trichoptera or Lepidoptera.
Hymenoptera included a similar spectrum of sawflies (Rasnitsyn 1968) and parasitoid lineages that were more diverse than those in the Yanliao Biota. The hymenopteran
parasitoid lineages include the parasitoid sawfly Orussoidea (
†
Karatavitidae), and
parasitoid wasps Ephialtitoidea (
†
Ephialtitidae), Megalyroidea (Megalyridae),
Evanioidea (
†
Anomopterellidae,
†
Praeaulacidae), Proctotrupoidea (Heloridae,
†
Mesoserphidae) and the basal aculeate lineage Chrysidoidea (
†
“Bethylonymidae”)
(Kozlov 1968; Rasnitsyn 2002) (Table 11.1). The same number of eight, familylevel lineages of hymenopteran parasitoids was present in the Karatau Biota as in
the Yanliao Biota, but with two family substitutions and a doubling of species to
approximately 30, indicating evolutionary stability at the family level but an
increased proliferation of species. In general, it appears that parasitoids of the
Karatau Biota represent continued stability of major parasitoid lineages when compared with the Yanliao Biota, but increased generation of genera and species at
lower taxonomic levels.
The third insect fauna considered during the initial expansion of the MMPR
comes from the Lulworth Formation, constituting the lower unit of the Purbeck
Limestone Group, and its upper unit, the Durlston Formation, which crop out along
the Vale of Wardour area in Dorset, southern England (Rasnitsyn et al. 1998; Coram
et al. 2000; Coram and Jepson 2012). The age of the Purbeck strata is earliest
Cretaceous (Berriasian Stage), with a corresponding age date of 145–139 Ma
(Walker et al. 2013), and is approximately 16 million years younger than the Karatau
Biota. The strata of the Purbeck Biota containing the insect fossils consist of finegrained, mostly thinly bedded limestone occasionally interrupted by algal mat layers (Coram 2003). Parasitoid-containing lineages of Neuroptera, Coleoptera,
Strepsiptera, Diptera, Trichoptera, and Lepidoptera are absent. However, as in the
preceding two fossil units, a near full complement of sawflies is present, excluding
the Orussoidea—the only parasitoid sawfly clade. Apocritan wasps consist of
Megalyroidea (Megalyridae), Evanioidea (cf. Aulacidae,
†
Baissidae,
†
incertae
sedis),
Proctotrupoidea
(Diapriidae,
Proctotrupidae),
Ichneumonoidea
(Ichneumonidae), the aculeate lineages Chrysidoidea (
†
“Bethylonymidae”) and
Apoidea (Sphecidae), and the undetermined apocritan Apocrites (Coram and Jepson
2012) (Table 11.1). Including unassigned but distinctive lineages, there are 10
family- level lineages of hymenopteran parasitoids representing about 20 species in
the Purbeck Biota.
The Yixian Formation is the fourth insect fauna that highlights Phase 2 of the
MMPR (Fig. 11.19). Fossils from the Yixian Formation formed the distinctive Jehol
Biota that originate from seven or eight major, fossil-yielding localities, mostly in
Liaoning Province of northeastern China, west of Beipiao City and near Liaodong
Bay, an arm of the Yellow Sea (Zhang et al. 2010). Although once controversial
(Ren et al. 2010b), the date currently is established as mid Early Cretaceous, from
126 to 122 Ma (Swisher III et al. 1999), equivalent to late Barremian to early Aptian
in age (Walker et al. 2013), and approximately 18 million years younger than the
Purbeck Biota. The fossils occur in broad outcrops of lake deposits with lightcolored, highly oxidized, fine-grained mudstone and siltstone strata of buff-colored
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution