400
associated with complex ant nests. These two species occurred in the same, large,
amber pieces as the ant nests, and likely were myrmecophiles (Vršanský et al.
2019). (A myrmecophile is a commensal or parasite of the nest of an ant species.)
The authors interpret the two cockroach myrmecophile species as engaged in commensalistic and parasitic relationships with their ant hosts, consistent with morphological features in extant ant myrmecophiles. The significance of these extinct,
social parasitic cockroaches was their impact on the evolution of complex ant nests
in late Mesozoic tropical forests soon after the earliest evidence for ant eusociality
in the fossil record.
11.5.1.2 Dermaptera (Earwigs)
Modern Dermaptera (earwigs) consist of 3 suborders, 11 families and 203 genera,
and are nocturnal, hemimetabolous insects with distinctively short forewings and
terminal abdominal cerci modified into forceps-like pincers (Haas 2018). Two dermapteran lineages, Arixeniidae (bat earwigs) and Hemimeridae (rodent earwigs),
often are placed in their own suborders, and live ectoparasitically on mammals
(Popham 1984), although recent evidence suggests that the relationships may be
more commensalistic than parasitic (Haas 2018). Hemimeridae occur in subSaharan Africa on hamster rats and possess a suite of structures, such as dorsoventrally flattened bodies; short, grooved legs to cling to fur; and mouthparts for
abrading and feeding on host skin and surface fungi (Rehn and Rehn 1935, 1937;
Ashford 1970; Nakata and Maa 1974). Arixeniidae occur with molossid bats in
caves in Southeast Asia, and possess a matted pubescent body, long antennae and
long legs for rapid movement (Nakata and Maa 1974). A phylogenetic study of
Dermaptera (Kocarek et al. 2013) indicates that the Hemimeridae and Arixeniidae
each has a sister-group relationship with another dermapteran family, indicating that
the parasite lineages are convergently evolved, highly modified earwigs (Fig. 11.5).
These relationships indicate the potential for rapid change of parasite external morphology when a new trophic niche becomes available (Kocarek et al. 2013), likely
during the Eocene when their rodent and bat hosts initially diversified.
11.5.1.3 Psocoptera (Booklice, Psocids)
The term, Psocoptera (booklice, psocids) currently is not widely used, which is
partly attributable to the paraphyletic status that this ancient group of hemimetabolous insects has with respect to the nested clade of Phthiraptera, the parasitic lice
(Nagler and Haug 2015; Mockford 2018). Nevertheless, Psocoptera as used in this
report is a convenient reference to all intervening, free-living lineages of bark lice
that form the broader clade. The Psocoptera includes the variously ranked
Trogiomorpha, Amphientometae, Sphaeropsocidae, Psocomorpha (the overwhelming bulk of taxonomic diversity), and the book lice, Liposcelididae (Johnson et al.
2018). Of these, particular attention should be devoted to Liposcelididae. The
C. C. Labandeira and L. Li
associated with complex ant nests. These two species occurred in the same, large,
amber pieces as the ant nests, and likely were myrmecophiles (Vršanský et al.
2019). (A myrmecophile is a commensal or parasite of the nest of an ant species.)
The authors interpret the two cockroach myrmecophile species as engaged in commensalistic and parasitic relationships with their ant hosts, consistent with morphological features in extant ant myrmecophiles. The significance of these extinct,
social parasitic cockroaches was their impact on the evolution of complex ant nests
in late Mesozoic tropical forests soon after the earliest evidence for ant eusociality
in the fossil record.
11.5.1.2 Dermaptera (Earwigs)
Modern Dermaptera (earwigs) consist of 3 suborders, 11 families and 203 genera,
and are nocturnal, hemimetabolous insects with distinctively short forewings and
terminal abdominal cerci modified into forceps-like pincers (Haas 2018). Two dermapteran lineages, Arixeniidae (bat earwigs) and Hemimeridae (rodent earwigs),
often are placed in their own suborders, and live ectoparasitically on mammals
(Popham 1984), although recent evidence suggests that the relationships may be
more commensalistic than parasitic (Haas 2018). Hemimeridae occur in subSaharan Africa on hamster rats and possess a suite of structures, such as dorsoventrally flattened bodies; short, grooved legs to cling to fur; and mouthparts for
abrading and feeding on host skin and surface fungi (Rehn and Rehn 1935, 1937;
Ashford 1970; Nakata and Maa 1974). Arixeniidae occur with molossid bats in
caves in Southeast Asia, and possess a matted pubescent body, long antennae and
long legs for rapid movement (Nakata and Maa 1974). A phylogenetic study of
Dermaptera (Kocarek et al. 2013) indicates that the Hemimeridae and Arixeniidae
each has a sister-group relationship with another dermapteran family, indicating that
the parasite lineages are convergently evolved, highly modified earwigs (Fig. 11.5).
These relationships indicate the potential for rapid change of parasite external morphology when a new trophic niche becomes available (Kocarek et al. 2013), likely
during the Eocene when their rodent and bat hosts initially diversified.
11.5.1.3 Psocoptera (Booklice, Psocids)
The term, Psocoptera (booklice, psocids) currently is not widely used, which is
partly attributable to the paraphyletic status that this ancient group of hemimetabolous insects has with respect to the nested clade of Phthiraptera, the parasitic lice
(Nagler and Haug 2015; Mockford 2018). Nevertheless, Psocoptera as used in this
report is a convenient reference to all intervening, free-living lineages of bark lice
that form the broader clade. The Psocoptera includes the variously ranked
Trogiomorpha, Amphientometae, Sphaeropsocidae, Psocomorpha (the overwhelming bulk of taxonomic diversity), and the book lice, Liposcelididae (Johnson et al.
2018). Of these, particular attention should be devoted to Liposcelididae. The
C. C. Labandeira and L. Li