413
constitute about 32 modern genera of brachycerous flies that bite vertebrates using
a slashing and cutting strategy to obtain blood and fluids that pool up within lesions,
feeding strategy known as telmophagy (Bouchet and Lavaud 1999). Although the
earliest Tabanidae originate from the latest Middle Jurassic of northeastern China
(Ren 1998; Labandeira 2010), their original diet has been a mystery as to whether
they consumed nectar-like pollination drops or blood (Grimaldi and Engel 2005a).
This uncertainty arises because tabanids occur in the fossil record about 40 million
years before the appearance of angiosperms, although a broad variety of vertebrates
were present at this time. One hypothesis is that pangionine tabanids possessed
siphonate mouthparts and were consumers of pollination drops (Labandeira 2010),
whereas tabanine tabanids instead bore stylate mouthparts and were consumers of
vertebrate blood (Martins-Neto 2003; Lukashevich and Mostovski 2003; Labandeira
et al. 2007a, b). A less likely, alternative hypothesis is that for species of midMesozoic tabanids, males consumed plant fluids of ovulate organs in a pollination
mutualism. By contrast, conspecific females consumed vertebrate blood necessary
for oögenesis and indicative of parasitism, a feeding behavior that occurs in some
extant, hematophagous flies (Labandeira 2005).
Tabanids are known from younger earliest Cretaceous deposits of Brazil, the
United Kingdom and Russia, some with probable hematophagous habits (MartinsNeto 2003; Lukashevich and Mostovski 2003). One particular specimen from the
early Cretaceous of Transbaikalia is of large size with a robust body and gracile
legs, a head having an inflated clypeus and elongate proboscis encompassing hardened stylets, and antennae and palps that bear numerous, multiple types of sensillae
(Mostovski et al. 2003). These features are consistent with modern blood-feeding
tabanids (Lukashevich and Mostovski 2003). In Paleogene deposits, such as Baltic
Amber, the diversity of blood-feeding tabanids can be substantial, consisting contemporaneously of four genera and seven species of Chrysopinae and one genus and
species of Tabaninae (Pielowska et al. 2018).
Athericidae are a modestly speciose family of 13 genera in which adults feed on
nectar or blood and the aquatic larvae are predatory (Stuckenberg 1973). The athericid fossil record is unspectacular, with the exception of its earliest occurrence, the
unusual Qiyia jurassica from the Jiulongshan Formation of northeastern China
(Chen et al. 2014). Qiya jurassica possesses the most atypical attachment mechanisms for any ectoparasite: a thoracic sucker (Fig. 11.9). The aquatic fly larva
Q. jurassica is a stem-group athericid from the latest Middle Jurassic of northwestern China (Chen et al. 2014). The larva is interpreted to have inhabited streams with
rapid currents. In addition to this broad, ventral, circular attachment device occupying all three thoracic segments, Q. jurassica possesses a diminutive head with distinct piercing-and-sucking mouthparts, and stump-like abdominal prolegs for both
attachment and movement. The authors (Chen et al. 2014) suggest that feeding
occurred on salamander blood, although recent aquatic insect larvae in rapidly flowing streams with suction attachment discs are frequently predatory rather than parasitic in feeding habits (Leung 2017).
Rhagionoidea consist of one hematophagous family, Rhagionidae (snipe flies),
of medium- to large sized flies that contain about 16 genera, several of which are
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
constitute about 32 modern genera of brachycerous flies that bite vertebrates using
a slashing and cutting strategy to obtain blood and fluids that pool up within lesions,
feeding strategy known as telmophagy (Bouchet and Lavaud 1999). Although the
earliest Tabanidae originate from the latest Middle Jurassic of northeastern China
(Ren 1998; Labandeira 2010), their original diet has been a mystery as to whether
they consumed nectar-like pollination drops or blood (Grimaldi and Engel 2005a).
This uncertainty arises because tabanids occur in the fossil record about 40 million
years before the appearance of angiosperms, although a broad variety of vertebrates
were present at this time. One hypothesis is that pangionine tabanids possessed
siphonate mouthparts and were consumers of pollination drops (Labandeira 2010),
whereas tabanine tabanids instead bore stylate mouthparts and were consumers of
vertebrate blood (Martins-Neto 2003; Lukashevich and Mostovski 2003; Labandeira
et al. 2007a, b). A less likely, alternative hypothesis is that for species of midMesozoic tabanids, males consumed plant fluids of ovulate organs in a pollination
mutualism. By contrast, conspecific females consumed vertebrate blood necessary
for oögenesis and indicative of parasitism, a feeding behavior that occurs in some
extant, hematophagous flies (Labandeira 2005).
Tabanids are known from younger earliest Cretaceous deposits of Brazil, the
United Kingdom and Russia, some with probable hematophagous habits (MartinsNeto 2003; Lukashevich and Mostovski 2003). One particular specimen from the
early Cretaceous of Transbaikalia is of large size with a robust body and gracile
legs, a head having an inflated clypeus and elongate proboscis encompassing hardened stylets, and antennae and palps that bear numerous, multiple types of sensillae
(Mostovski et al. 2003). These features are consistent with modern blood-feeding
tabanids (Lukashevich and Mostovski 2003). In Paleogene deposits, such as Baltic
Amber, the diversity of blood-feeding tabanids can be substantial, consisting contemporaneously of four genera and seven species of Chrysopinae and one genus and
species of Tabaninae (Pielowska et al. 2018).
Athericidae are a modestly speciose family of 13 genera in which adults feed on
nectar or blood and the aquatic larvae are predatory (Stuckenberg 1973). The athericid fossil record is unspectacular, with the exception of its earliest occurrence, the
unusual Qiyia jurassica from the Jiulongshan Formation of northeastern China
(Chen et al. 2014). Qiya jurassica possesses the most atypical attachment mechanisms for any ectoparasite: a thoracic sucker (Fig. 11.9). The aquatic fly larva
Q. jurassica is a stem-group athericid from the latest Middle Jurassic of northwestern China (Chen et al. 2014). The larva is interpreted to have inhabited streams with
rapid currents. In addition to this broad, ventral, circular attachment device occupying all three thoracic segments, Q. jurassica possesses a diminutive head with distinct piercing-and-sucking mouthparts, and stump-like abdominal prolegs for both
attachment and movement. The authors (Chen et al. 2014) suggest that feeding
occurred on salamander blood, although recent aquatic insect larvae in rapidly flowing streams with suction attachment discs are frequently predatory rather than parasitic in feeding habits (Leung 2017).
Rhagionoidea consist of one hematophagous family, Rhagionidae (snipe flies),
of medium- to large sized flies that contain about 16 genera, several of which are
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution