389
2018). The cleptoparasite was identified based on distinctive features of its wing
that are affiliated with modern cleptoparasite bees. Similarly, modern Strepsiptera
(twisted-wing parasites) are endoparasitoids that have distinctive features, including
extreme sexual dimorphism between males and females and high levels of morphological specialization between immatures and adults (Askew 1971; Kathirithamby
2009). Three species of fossil Strepsiptera were found in 21 million-year-old
Dominican Amber (Kathirithamby and Grimaldi 1993) that display the same distinctive features as their extant strepsipteran relatives, indicating that the same parasitoid relationships have survived approximately 21 million years.
Amber provides a wealth of morphological, and indirectly behavioral, detail
occasionally revealing the consuming predator and its consumed prey item, often in
flagrante delicto (Labandeira 2014a). Examples include a dance fly with a nonbiting
midge clutched by its forelegs and an insect larva consuming a scuttle fly’s head
(Grimaldi 1996; Janzen 2002). As for parasitism, there are several occurrences of
modern-aspect Paleogene fleas attributable to a modern clade (Poinar Jr 2015).
Older, Late Jurassic to Early Cretaceous stem-group lineages have been attributed
to modern Siphonaptera (Gao et al. 2012, 2014) based on several common, structural features (but see Dittmar et al. 2016; Leung 2021). Mesozoic evidence for
parasitoids also occurs in amber, but is qualitatively different by providing more
detail of external insect structure (Labandeira 2014a). Poinar (2013) shows several
such examples of parasitoids that involve insects.
11.3.3 Structural and Functional Attributes
Structural evidence historically has been the standard mode for inferring the presence of parasitism or parasitoidism in the fossil record. Many examples illustrate
body structures, especially mouthparts, attachment devices and ovipositors, of a
particular larva or adult that indicate a parasite or parasitoid life habit during the
later Mesozoic and Cenozoic (Leung 2017). Body structures strongly implicating
parasitoidism include those from Coleoptera (Engel 2005a; Poinar Jr 2009), Diptera
(Rocha et al. 2015; Zhang et al. 2016) and Hymenoptera (Barling et al. 2013;
Spasojevic et al. 2017). Of special note is synchrotron X-ray microtomography
(Labandeira 2014a), that has been used for documenting parasitoid interactions
with their hosts caught in the act (van de Kamp et al. 2018). In that study (van de
Kamp et al. 2018) 55 parasitoidization events were three-dimensionally imaged that
recorded four species of hymenopteran endoparasitoids in a dipteran pupal host.
The life habits of predators, parasites and parasitoids frequently can be deduced
from their functional morphology (Leung 2017). The fossil record provides considerable evidence for predation, such as silk-producing spigots in a Devonian insectivorous spider likely used for trapping of prey items (Fig. 11.1b). The fossil record
also provides evidence, such as wing eyespots of a cnemidolestid archaeorthopteran
(Fig. 11.1d), evidently a potential prey item, to deflect attention and avoid
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
2018). The cleptoparasite was identified based on distinctive features of its wing
that are affiliated with modern cleptoparasite bees. Similarly, modern Strepsiptera
(twisted-wing parasites) are endoparasitoids that have distinctive features, including
extreme sexual dimorphism between males and females and high levels of morphological specialization between immatures and adults (Askew 1971; Kathirithamby
2009). Three species of fossil Strepsiptera were found in 21 million-year-old
Dominican Amber (Kathirithamby and Grimaldi 1993) that display the same distinctive features as their extant strepsipteran relatives, indicating that the same parasitoid relationships have survived approximately 21 million years.
Amber provides a wealth of morphological, and indirectly behavioral, detail
occasionally revealing the consuming predator and its consumed prey item, often in
flagrante delicto (Labandeira 2014a). Examples include a dance fly with a nonbiting
midge clutched by its forelegs and an insect larva consuming a scuttle fly’s head
(Grimaldi 1996; Janzen 2002). As for parasitism, there are several occurrences of
modern-aspect Paleogene fleas attributable to a modern clade (Poinar Jr 2015).
Older, Late Jurassic to Early Cretaceous stem-group lineages have been attributed
to modern Siphonaptera (Gao et al. 2012, 2014) based on several common, structural features (but see Dittmar et al. 2016; Leung 2021). Mesozoic evidence for
parasitoids also occurs in amber, but is qualitatively different by providing more
detail of external insect structure (Labandeira 2014a). Poinar (2013) shows several
such examples of parasitoids that involve insects.
11.3.3 Structural and Functional Attributes
Structural evidence historically has been the standard mode for inferring the presence of parasitism or parasitoidism in the fossil record. Many examples illustrate
body structures, especially mouthparts, attachment devices and ovipositors, of a
particular larva or adult that indicate a parasite or parasitoid life habit during the
later Mesozoic and Cenozoic (Leung 2017). Body structures strongly implicating
parasitoidism include those from Coleoptera (Engel 2005a; Poinar Jr 2009), Diptera
(Rocha et al. 2015; Zhang et al. 2016) and Hymenoptera (Barling et al. 2013;
Spasojevic et al. 2017). Of special note is synchrotron X-ray microtomography
(Labandeira 2014a), that has been used for documenting parasitoid interactions
with their hosts caught in the act (van de Kamp et al. 2018). In that study (van de
Kamp et al. 2018) 55 parasitoidization events were three-dimensionally imaged that
recorded four species of hymenopteran endoparasitoids in a dipteran pupal host.
The life habits of predators, parasites and parasitoids frequently can be deduced
from their functional morphology (Leung 2017). The fossil record provides considerable evidence for predation, such as silk-producing spigots in a Devonian insectivorous spider likely used for trapping of prey items (Fig. 11.1b). The fossil record
also provides evidence, such as wing eyespots of a cnemidolestid archaeorthopteran
(Fig. 11.1d), evidently a potential prey item, to deflect attention and avoid
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution