404
specimen’s life style was an ectoparasite of a warm-blooded vertebrate possessing
hair, such as a pterosaur, or possibly a mammal of large size (Rasnitsyn and Zherikhin
1999). Nevertheless, the specimen does reveal structures similar to chewing lice of the
Ishnocera such as a highly flattened body, a single clawed tarsus and robust truck
tracheae and spiracles, but its’ very large size would be inconsistent with parasitization of any host except a large vertebrate. However, contrary interpretations regarding
the affinities of the fossil suggest that this specimen is not a phthirapteran (Wappler
et al. 2004; Dalgleish et al. 2006). Major features indicating that the specimen is not a
phthirapteran is the absence or lack of preservation of mandibles and the presence of
large lateral head processes (Dalgleish et al. 2006).
From the Eckfeld maar crater beds in Germany, of middle Eocene (44 Ma) age,
is Megamenopon rasnitsyni (Amblycera: Menoponidae), a member of primitive
chewing lice (Wappler et al. 2004). This specimen exhibits similarities to amblyceran feather lice that parasitize Anseriformes (ducks, geese and swans) and
Charadiformes (shorebirds). The presence of feather chaff in gut contests of the
specimen confirms its life habit as an ectoparasite (Wappler et  al. 2004). From
nearby Baltic Amber, a deposit of approximately the same age as Eckfeld, Voigt
(1952) described phthirapteran eggs cemented to mammalian hair, likely belonging
to an unknown sucking louse of Anoplura.
11.5.1.5 Hemiptera (Bugs)
Within the Hemiptera, it is only the suborder Heteroptera (true bugs) and its constituent infraorder, Cimicomorpha, which contains parasitic lineages engaged in
hematophagy. Four cimicomorph lineages, Cimicidae (bed bugs), Polyctenidae (bat
bugs), Reduviidae (assassin bugs), and extinct Torirostratidae, contain the hematophagous genera within Hemiptera. Given established phylogenies within the
Cimicomorpha (Schuh et al. 2009; Weirauch and Munro 2009; also see Yao et al.
2014), hematophagous parasitism originated in the Hemiptera three times: once in
Reduviidae, once in the Polyctenidae  +  Cimicidae clade, and once in the extinct
Torirostratidae.
Cimicidae consists of 6 subfamilies and 21 genera, including the common bed
bug Cimex lectularis, which are obligate parasites of warm-blooded vertebrates,
including humans. As a group, cimicids are frequently narrow host specialists, and
rarely are vectors for disease-causing viruses and other pathogens (Reinhardt and
Siva-Jothy 2007). Cimicids are attracted to hosts by several cues, including temperature, carbon dioxide gradients and animal kairomones, and feed once every 3–7
days (Reinhardt and Siva-Jothy 2007).
Polyctenidae are a family of rare subtropical bugs, the probable sister-group of
Cimicidae, that consist of two subfamilies and five genera that are ectoparasites of
bats and exhibit substantial host specialization. The family derives its name from
the presence of prominent combs that cover several regions of the body, including
the antennal bases, gena of the head, and pronotum, prosternum, mesonotum, ventral surfaces of the abdominal segments and tarsi (Askew 1971). Members of
C. C. Labandeira and L. Li
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