417
increasing in size through several instar shifts up to 5 cm long, eventually maturing
and leaving the host during early Arctic spring and pupating on the ground, where
they emerge as adult flies after several weeks to complete the life cycle (Asbakk
et al. 2014). The cumulative load of reindeer warble fly infestation is particularly
debilitating on reindeer cows, particularly during calving season in late summer
(Ballesteros et al. 2011). Thematically, this life cycle of infestation is similar to bot
flies attacking a variety of other large mammals, especially bovids (e.g., Oryan
et al. 2009).
Calliphoridae are a diverse, worldwide family consisting of 75 genera, some of
which have larvae (screwworms) that cause myiasis and other diseases such as dysentery and paratuberculosis that especially affect livestock (Thomas and Mangan
1989) (Table 11.1). Calliphoridae are of vital importance in carrion communities,
assisting the degradation of animal tissues. Several occurrences (puparia) of
Calliphoridae are late Neogene in age, the oldest is upper Pliocene, from 3.6 to 2.6
million years ago (Kitching 1980). An earlier record of two puparia in an ironstone
nodule attributed to the Calliphoridae, from the late Cretaceous of Canada
(Campanian Stage) and described by McAlpine (1970), is erroneous and are attributed to Cyclorrhapha (Grimaldi and Engel 2005a). Since no calyptrate group is
known from the Mesozoic (Grimaldi and Engel 2005a), the calyptrate Calliphoridae
likely originated during the more recent Paleogene and was parasitic on mammals
of medium to large size.
A closely related lineage to the Calliphoridae is the Mesembrinellidae, consisting of three subfamilies and nine genera that occur in humid, primary forests of the
Neotropics. Some adults feed on blood and are parasites whereas other species are
found on decomposing animal material and fermented substances in fruits (Marinho
et al. 2017). The only fossil occurrence is Mesembrinella caenozoica from early
Miocene amber (Fig. 11.10), from the Dominican Republic (21 Ma). Their origin is
placed during the Eocene (Cerretti et al. 2017).
Mystacinobiidae is a closely related family to Calliphoridae, consisting of the
enigmatic, monotypic Mystacinobia zelandica (New Zealand bat fly) that recently
has been discovered (Gleeson et al. 2000). This group is a spider-like, wingless,
structurally highly modified fly that feeds on bat guano and is phoretic and possibly
parasitic on the New Zealand short-tailed bat (Holloway 1976).
Excluded from the list of parasitic Diptera, Strashilidae is a group of midMesozoic, eastern Eurasian insects exemplified by the morphologically bizarre
Strashila incredibilis (Fig. 11.1e; Rasnitsyn 1992). They possess features that
superficially resemble some ectoparasitic insects. These features include dorsoventral flattening; legs with highly swollen metafemora and grasping terminal claws;
prominent lateral, gill-like appendages of the abdomen; and a head with sunken
antennae and a ventrally directed, short beak housing reduced piercing-and- sucking
mouthparts (Huang et al. 2013b). Males were winged whereas females were wingless. Once thought as terrestrial ectoparasites of pterosaurs and possibly affiliated
with Siphonaptera (Rasnitsyn 1992), the group together with Tarwiniidae were considered as sister group to Saurophthiridae (Rasnitsyn and Quicke 2002; Nagler and
Haug 2015). Others proposed that Strashilidae as a lineage originating deeper
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
increasing in size through several instar shifts up to 5 cm long, eventually maturing
and leaving the host during early Arctic spring and pupating on the ground, where
they emerge as adult flies after several weeks to complete the life cycle (Asbakk
et al. 2014). The cumulative load of reindeer warble fly infestation is particularly
debilitating on reindeer cows, particularly during calving season in late summer
(Ballesteros et al. 2011). Thematically, this life cycle of infestation is similar to bot
flies attacking a variety of other large mammals, especially bovids (e.g., Oryan
et al. 2009).
Calliphoridae are a diverse, worldwide family consisting of 75 genera, some of
which have larvae (screwworms) that cause myiasis and other diseases such as dysentery and paratuberculosis that especially affect livestock (Thomas and Mangan
1989) (Table 11.1). Calliphoridae are of vital importance in carrion communities,
assisting the degradation of animal tissues. Several occurrences (puparia) of
Calliphoridae are late Neogene in age, the oldest is upper Pliocene, from 3.6 to 2.6
million years ago (Kitching 1980). An earlier record of two puparia in an ironstone
nodule attributed to the Calliphoridae, from the late Cretaceous of Canada
(Campanian Stage) and described by McAlpine (1970), is erroneous and are attributed to Cyclorrhapha (Grimaldi and Engel 2005a). Since no calyptrate group is
known from the Mesozoic (Grimaldi and Engel 2005a), the calyptrate Calliphoridae
likely originated during the more recent Paleogene and was parasitic on mammals
of medium to large size.
A closely related lineage to the Calliphoridae is the Mesembrinellidae, consisting of three subfamilies and nine genera that occur in humid, primary forests of the
Neotropics. Some adults feed on blood and are parasites whereas other species are
found on decomposing animal material and fermented substances in fruits (Marinho
et al. 2017). The only fossil occurrence is Mesembrinella caenozoica from early
Miocene amber (Fig. 11.10), from the Dominican Republic (21 Ma). Their origin is
placed during the Eocene (Cerretti et al. 2017).
Mystacinobiidae is a closely related family to Calliphoridae, consisting of the
enigmatic, monotypic Mystacinobia zelandica (New Zealand bat fly) that recently
has been discovered (Gleeson et al. 2000). This group is a spider-like, wingless,
structurally highly modified fly that feeds on bat guano and is phoretic and possibly
parasitic on the New Zealand short-tailed bat (Holloway 1976).
Excluded from the list of parasitic Diptera, Strashilidae is a group of midMesozoic, eastern Eurasian insects exemplified by the morphologically bizarre
Strashila incredibilis (Fig. 11.1e; Rasnitsyn 1992). They possess features that
superficially resemble some ectoparasitic insects. These features include dorsoventral flattening; legs with highly swollen metafemora and grasping terminal claws;
prominent lateral, gill-like appendages of the abdomen; and a head with sunken
antennae and a ventrally directed, short beak housing reduced piercing-and- sucking
mouthparts (Huang et al. 2013b). Males were winged whereas females were wingless. Once thought as terrestrial ectoparasites of pterosaurs and possibly affiliated
with Siphonaptera (Rasnitsyn 1992), the group together with Tarwiniidae were considered as sister group to Saurophthiridae (Rasnitsyn and Quicke 2002; Nagler and
Haug 2015). Others proposed that Strashilidae as a lineage originating deeper
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution