410
traits are important. For example, the labial palps form a short, segmented, robust
structure in Saurophthirus and modern fleas, whereas they form a long, nonsegmented, streamlined, more gracile structure enveloping the mouthpart stylets in
all other mid-Mesozoic flea-like lineages except Tarwinia (Gao et al. 2014). This
indicates that the Saurophthirus + modern fleas constitute an expanded Siphonaptera
clade that is distantly related to all other mid-Mesozoic fleas.
The earliest hosts for these early true flea lineages were mammals (Perrichot
et al. 2012), although four, separate, parasite shifts apparently colonized birds most
likely through introductions from ground nesting flea taxa (Whiting et al. 2008; Zhu
et al. 2015). During the Cenozoic, several lineages of fleas are documented in
ambers, although the most commonly encountered are common fleas (Pulicidae),
particularly in Baltic and Dominican ambers (Lewis and Grimaldi 1997; Perrichot
et al. 2012; Poinar Jr 2015). Rodent fleas of Ctenophthalmidae and Hystrichopsyllidae
(Fig. 11.1l) were described from Baltic Amber (Dampf 1910; Pielowska et al. 2018).
Marsupial fleas of Rhopalopsyllidae are known from Dominican Amber (Grimaldi
and Engel 2005a).
11.5.1.8 Diptera (Flies)
Diptera constitute the greatest variety of parasitism, principally through bloodfeeding (hematophagy), of any insect group, and are responsible for the widespread
vectoring of parasites that often produce blood diseases principally to vertebrates,
including humans. Eight major clades encompass 15 families that collectively are
responsible for almost all of the documented parasitic relationships between Diptera
and their vertebrate and other hosts (Lehane 1991; Lukashevich and Mostovski
2003). At the family level, hematophagy originated minimally 12 times (Wiegmann
et al. 2011), although at lower taxonomic levels of origination are much greater.
Dipteran hematophagous clades often are assembled into a nematocerous group
(Culicoidea, Chironomoidea, Psychodiodea), a brachycerous group (Tabanoidea,
Rhagionoidea), and a cyclorrhaphous group (Muscoidea, Hippoboscoidea,
Oestroidea). Notably, these clades collectively represent the widest variety of
mouthpart types in insects (Labandeira 1997, 2019), and house a diversity of feeding styles. Most of these families have members that are vectors of a variety of
infectious diseases (Lehane 1991), and apparently opportunities were seized in
occupation of new ecological niches for transmission of blood parasites sometime
during the Early Cretaceous (Leung 2017). The pathogen fossil record indicates that
such disease transmission commenced during the Early Cretaceous and has
expanded to the Recent (Labandeira 2014a; Poinar Jr 2018, 2021).
The Culicoidea is a major clade of nematocerous flies (Wiegmann et al. 2011)
that evolved lineages parasitic on vertebrates through blood feeding. One of the
earliest occurrences of Culicoidea is from Late Triassic (late Carnian) strata of the
Solite Quarry in southernmost Virginia, U.S.A. (Blagoderov et al. 2007). A particular specimen (Fig. 11.8), unassignable to family, displays the anterior thoracic
region, head capsule, eyes, and sufficiently well preserved mouthparts to determine
that the proboscis likely has a design consistent with feeding on pollination drops or
C. C. Labandeira and L. Li
traits are important. For example, the labial palps form a short, segmented, robust
structure in Saurophthirus and modern fleas, whereas they form a long, nonsegmented, streamlined, more gracile structure enveloping the mouthpart stylets in
all other mid-Mesozoic flea-like lineages except Tarwinia (Gao et al. 2014). This
indicates that the Saurophthirus + modern fleas constitute an expanded Siphonaptera
clade that is distantly related to all other mid-Mesozoic fleas.
The earliest hosts for these early true flea lineages were mammals (Perrichot
et al. 2012), although four, separate, parasite shifts apparently colonized birds most
likely through introductions from ground nesting flea taxa (Whiting et al. 2008; Zhu
et al. 2015). During the Cenozoic, several lineages of fleas are documented in
ambers, although the most commonly encountered are common fleas (Pulicidae),
particularly in Baltic and Dominican ambers (Lewis and Grimaldi 1997; Perrichot
et al. 2012; Poinar Jr 2015). Rodent fleas of Ctenophthalmidae and Hystrichopsyllidae
(Fig. 11.1l) were described from Baltic Amber (Dampf 1910; Pielowska et al. 2018).
Marsupial fleas of Rhopalopsyllidae are known from Dominican Amber (Grimaldi
and Engel 2005a).
11.5.1.8 Diptera (Flies)
Diptera constitute the greatest variety of parasitism, principally through bloodfeeding (hematophagy), of any insect group, and are responsible for the widespread
vectoring of parasites that often produce blood diseases principally to vertebrates,
including humans. Eight major clades encompass 15 families that collectively are
responsible for almost all of the documented parasitic relationships between Diptera
and their vertebrate and other hosts (Lehane 1991; Lukashevich and Mostovski
2003). At the family level, hematophagy originated minimally 12 times (Wiegmann
et al. 2011), although at lower taxonomic levels of origination are much greater.
Dipteran hematophagous clades often are assembled into a nematocerous group
(Culicoidea, Chironomoidea, Psychodiodea), a brachycerous group (Tabanoidea,
Rhagionoidea), and a cyclorrhaphous group (Muscoidea, Hippoboscoidea,
Oestroidea). Notably, these clades collectively represent the widest variety of
mouthpart types in insects (Labandeira 1997, 2019), and house a diversity of feeding styles. Most of these families have members that are vectors of a variety of
infectious diseases (Lehane 1991), and apparently opportunities were seized in
occupation of new ecological niches for transmission of blood parasites sometime
during the Early Cretaceous (Leung 2017). The pathogen fossil record indicates that
such disease transmission commenced during the Early Cretaceous and has
expanded to the Recent (Labandeira 2014a; Poinar Jr 2018, 2021).
The Culicoidea is a major clade of nematocerous flies (Wiegmann et al. 2011)
that evolved lineages parasitic on vertebrates through blood feeding. One of the
earliest occurrences of Culicoidea is from Late Triassic (late Carnian) strata of the
Solite Quarry in southernmost Virginia, U.S.A. (Blagoderov et al. 2007). A particular specimen (Fig. 11.8), unassignable to family, displays the anterior thoracic
region, head capsule, eyes, and sufficiently well preserved mouthparts to determine
that the proboscis likely has a design consistent with feeding on pollination drops or
C. C. Labandeira and L. Li