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a 46 million-year-old Eocene mosquito from the Kishenehn Formation of northwestern Montana with blood in its distended abdomen (Greenwalt 2021; Greenwalt
et al. 2013). Heme-derived porphyrin molecules of hemoglobin were detected and
molecularly characterized by advanced mass spectrometry and other techniques,
indicating the presence of blood feeding, although the source of the vertebrate blood
remains unknown.
The Chironomoidea include the principal blood-feeding groups of
Ceratopogonidae (biting midges, no-see-ums) and Simuliidae (black flies, buffalo
gnats). While extant Chironomidae (non-biting midges) are considered as predominately feeding on pollen, nectar, honeydew and other carbohydrate-rich food, extinct
forms were hematophagous based on mouthpart structure (Azar and Nel 2012).
Chironomidae extend to the Late Triassic–Early Jurassic boundary interval
(Grimaldi and Engel 2005a), making them among the earliest hematophagous
Diptera. Ceratopogonidae are represented by several, prominent, extant genera that
are blood feeders, and the family has a good fossil record (Borkent 2001), extending
to the Early Cretaceous (Berriasian Stage) at about 140  Ma. There are several
instances of ceratopogonid parasitism in the amber fossil record. One example is a
ceratopogonid specimen from Myanmar Amber (99  Ma) that hosted in its gut
oocysts of the malarial parasite, Paleohaemoproteus burmacis, a pathogen similar
to modern species found in reptiles and birds (Poinar Jr and Telford Jr 2005), and
likely representing an ancient association. From the same deposit, a second ceratopogonid specimen, Leptoconops nosopheris, vectored the trypanosome
Paleotrypanosoma burmanicus that may have produced trypanosomiasis-like symptoms in an unknown vertebrate host (Poinar 2021).
The fossil record of the Simuliidae begins during the Late Jurassic (Oxfordian
Stage) and has a fair record of occurrences up to the recent (Currie and Grimaldi
2000). However, black fly–parasite associations are rare. The earliest fossil Simuliidae
has been described from Kazakhstan (Kalugina 1991). From these compressionimpression deposits, it appears that the blood-feeding habit in Simuliidae originated
early within the clade, and may be the ancestral diet (Kalugina 1991).
The Psychodoidea is a more ancient nematocerous lineage than Chironomoidea
that extends to the Late Triassic (Fraser et al. 1996). The only blood-feeding lineage
of Psychodoidea is Psychodidae (moth flies), of which the subfamily Phlebotominae
(sand flies) represent the only origin of hematophagy within the larger clade
(Grimaldi and Engel 2005a, b) and are a major cause globally for transmission of
several modern diseases in animals (Lehane 1991). One Cretaceous association
involves the phlebotomine Palaeomyia burmitis that vectored a trypanosome parasite, Paleoleishmania proterus, which presumably developed in reptilian blood cells
consumed by the sand fly (Poinar Jr 2004a; Poinar Jr and Poinar 2004a, b). This
sand fly–trypanosome–reptile parasitic interaction occurs in mid Cretaceous
Myanmar amber (Poinar 2021), and suggests a pathological relationship similar to
modern leishmaniasis, a debilitating disease creating ulcers of the skin and viscera
in vertebrates.
Tabanoidea consist of two families, Tabanidae (horse flies, deer flies) and
Athericidae (watersnipe flies) that are involved in blood feeding. Tabanidae
C. C. Labandeira and L. Li
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