458
Kathirithamby 2009). It appears that the early hosts of Strepsiptera may have been
more peripheral to their current host distribution, such as wood frequenting
cockroaches.
11.5.2.4 Diptera (Flies)
Dipteran parasitoids represent a much wider spectrum of hosts than does their other
major parasitoid competitor, Hymenoptera. Dipteran parasitoids attack 22 orders of
organisms within the five phyla of Platyhelminthes (flatworms), Mollusca (aquatic
and terrestrial gastropods), Annelida (oligochaetes), Arthropoda (including millipedes, crustaceans and arachnids) and Chordata (amphibians) (Table 11.1) (Morris
1998). This host spectrum represents the dominant terrestrial animal phyla, except
for Nematoda. Diptera have acquired the parasitoid habit minimally 21 times
(Eggleton and Belshaw 1992), representing about 16,000 species or 20% of all dipteran species (Feener and Brown 1997), although Weinstein and Kuris (2016) list 60
times after an exhaustive literature search. The actual number of instances of dipteran parasitoidism will be shown to be about 100 times, once the life habits and
phylogenetic relationships of more obscure lineages, particularly the diverse
Asilidae, Phoridae and Sarcophagidae, are known (Eggleton and Belshaw 1992).
Notably, in three of the four most diverse families—Bombyliidae, Conopidae and
Tachinidae—parasitoidism has evolved only once (Feener and Brown 1997; Yeates
and Greathead 1997).
Nine major dipteran lineages at the superfamily level harbor parasitoids.
Culicomorpha consists of Chironomidae, some of which are ectoparasitoids of pulmonate gastropod immatures and mayfly naiads. Other minor origins of parasitoidism in Diptera include a single origin from microphagy in Chironomidae (nonbiting
midges), and another single origin from mycophagy in Mycetophilidae (fungus
gnats). Although Chironomidae extends to the Late Triassic, it is unlikely that these
associations are as old. Bibionomorpha (e.g., Poinar Jr 2010) include endoparasitoids of flatworms and the nymphs and adults of sternorrhynchan hemipterans,
whose associations most likely are relatively recent and Neogene in origin (Eggleton
and Belshaw 1992). Nemestrinimorpha are obligate endoparasitoids of spiders and
mites, as well as grasshoppers and beetles, and collectively attack all host life stages.
Host associations of Nemestrinimorpha likely extend to the Late Triassic for tanglevein flies (Nemestrinidae). The spider flies (Acroceridae) are a more recently
evolved lineage that range from the Late Jurassic to the recent (Mostovski 1998).
They are endoparasitoids of spiders and mites and are particularly associated with
the parasitoidization of several major clades of araneomorph spiders (Gillung et al.
2018; Fig. 11.14). This lineage originated during the Middle Jurassic and diversified
during the Late Cretaceous to Paleogene. The Archisargoidea is the only extinct
dipteran superfamily that contains a parasitoid lineage (Fig. 11.15), the highly probable endoparasitoid clade Eremochaetidae, known from the Late Jurassic to earliest
Late Cretaceous (Grimaldi and Barden 2016). Although considered typically predaceous (Fig. 11.1f), Asilomorpha also are ectoparasitoids or endoparasitoids,
C. C. Labandeira and L. Li
Kathirithamby 2009). It appears that the early hosts of Strepsiptera may have been
more peripheral to their current host distribution, such as wood frequenting
cockroaches.
11.5.2.4 Diptera (Flies)
Dipteran parasitoids represent a much wider spectrum of hosts than does their other
major parasitoid competitor, Hymenoptera. Dipteran parasitoids attack 22 orders of
organisms within the five phyla of Platyhelminthes (flatworms), Mollusca (aquatic
and terrestrial gastropods), Annelida (oligochaetes), Arthropoda (including millipedes, crustaceans and arachnids) and Chordata (amphibians) (Table 11.1) (Morris
1998). This host spectrum represents the dominant terrestrial animal phyla, except
for Nematoda. Diptera have acquired the parasitoid habit minimally 21 times
(Eggleton and Belshaw 1992), representing about 16,000 species or 20% of all dipteran species (Feener and Brown 1997), although Weinstein and Kuris (2016) list 60
times after an exhaustive literature search. The actual number of instances of dipteran parasitoidism will be shown to be about 100 times, once the life habits and
phylogenetic relationships of more obscure lineages, particularly the diverse
Asilidae, Phoridae and Sarcophagidae, are known (Eggleton and Belshaw 1992).
Notably, in three of the four most diverse families—Bombyliidae, Conopidae and
Tachinidae—parasitoidism has evolved only once (Feener and Brown 1997; Yeates
and Greathead 1997).
Nine major dipteran lineages at the superfamily level harbor parasitoids.
Culicomorpha consists of Chironomidae, some of which are ectoparasitoids of pulmonate gastropod immatures and mayfly naiads. Other minor origins of parasitoidism in Diptera include a single origin from microphagy in Chironomidae (nonbiting
midges), and another single origin from mycophagy in Mycetophilidae (fungus
gnats). Although Chironomidae extends to the Late Triassic, it is unlikely that these
associations are as old. Bibionomorpha (e.g., Poinar Jr 2010) include endoparasitoids of flatworms and the nymphs and adults of sternorrhynchan hemipterans,
whose associations most likely are relatively recent and Neogene in origin (Eggleton
and Belshaw 1992). Nemestrinimorpha are obligate endoparasitoids of spiders and
mites, as well as grasshoppers and beetles, and collectively attack all host life stages.
Host associations of Nemestrinimorpha likely extend to the Late Triassic for tanglevein flies (Nemestrinidae). The spider flies (Acroceridae) are a more recently
evolved lineage that range from the Late Jurassic to the recent (Mostovski 1998).
They are endoparasitoids of spiders and mites and are particularly associated with
the parasitoidization of several major clades of araneomorph spiders (Gillung et al.
2018; Fig. 11.14). This lineage originated during the Middle Jurassic and diversified
during the Late Cretaceous to Paleogene. The Archisargoidea is the only extinct
dipteran superfamily that contains a parasitoid lineage (Fig. 11.15), the highly probable endoparasitoid clade Eremochaetidae, known from the Late Jurassic to earliest
Late Cretaceous (Grimaldi and Barden 2016). Although considered typically predaceous (Fig. 11.1f), Asilomorpha also are ectoparasitoids or endoparasitoids,
C. C. Labandeira and L. Li