416
Germany, of late Oligocene age (Chattian Stage) at approximately 25.5 Ma (Statz
1940; Maa 1966). Maa (1966) concluded that the specimen was an early, basal lineage of Hippoboscidae. A more recent find of a Hippoboscid is from the latest
Miocene Messinian Stage, approximately 6 million years in age (Bradley and
Landini 1984). However, these occurrences say little about the evolutionary biology
of hippoboscids (Dittmar et al. 2006). An alternative method is a molecular phylogenetic approach (Dittmar et al. 2006) that places the Hippoboscidae as the sistergroup to the Glossinidae, with an origin approximately at the same time as the
separation of the Nycteribiidae and Streblidae during the late Eocene (De Moya 2019).
Nycteribiidae (bat flies) are 12 genera of dorsoventrally flattened, spider-like
flies lacking eyes and wings that are obligate, host specific parasites on bats.
Nycteribiids have been implicated in the transmission of bat malaria in Africa
(Obame-Nkoghe et al. 2016). They lack a fossil record. Related to Nycteribiidae are
Streblidae (bat flies), consisting of 5 subfamilies and approximately 33 genera of
blood-feeding flies that are ectoparasitic specialists on a variety of bats (Lehane
1991). Some genera are involved in transmission of bat malaria (Obame-Nkoghe
et al. 2016). Although the oldest streblid is early Miocene in age (Poinar Jr and
Brown 2012), the clade is thought to have originated during the Eocene, based on
the fossil record of the closely related Glossinidae (Dittmar et al. 2006). From the
same early Miocene deposit, an unaffiliated streblid harbored Vetufebrus ovatus
(Haemospororida: Plasmodiidae), a bat malarial parasite (Poinar Jr 2011) that
apparently is not vectored by any known extant streblid (Leung 2017). This suggests that this particular parasitic relationship of malaria may be a disease that has
become extinct for streblid vectors and their bat hosts (compare van Dijk and De
Baets 2021 for other putative examples).
Oestroidea contains four families, Oestridae (bot flies), Calliphoridae (blow flies,
carrion flies, greenbottles), Mesembrinellidae (blow flies) and Mystacinobiidae
(New Zealand bat fly), which contain significant numbers of parasitic, bloodfeeding species. Parasitism likely originated five times in the Oestroidea, based on
genus-level phylogenies (Pape 2001; Kutty et al. 2001). As cyclorrhaphan calyptrates, Oestroidea are a recent clade that originated during the late Paleocene to
early Eocene epochs of the Cenozoic, about 60–40 million years ago (Cerretti et al.
2017). The most iconic of the oestroid parasitic families, Oestridae consist of about
160 species (Wood 2006) and contain the most conspicuous, diverse and noxious of
parasites, as they are internal parasites of mammals and frequently inhabit their
host’s flesh, occasionally using an intermediate vector such as a dipteran to complete their life cycle (Colwell et al. 2006). A typical life cycle is illustrated by the
reindeer warble fly, Hypoderma tarandi, documented on reindeer, Rangifer tarandus. The fly initially lays eggs during Arctic summer at the base of thin hairs adjacent the skin, followed by hatching and then penetration of the first-instar larva into
the skin, soon resulting in development of nodules (warbles) under the skin while
the host’s proteins from subcutaneous tissue are ingested by the larva (Asbakk et al.
2014). During the following Arctic winter, the larva continues to consume proteins,
C. C. Labandeira and L. Li
Germany, of late Oligocene age (Chattian Stage) at approximately 25.5 Ma (Statz
1940; Maa 1966). Maa (1966) concluded that the specimen was an early, basal lineage of Hippoboscidae. A more recent find of a Hippoboscid is from the latest
Miocene Messinian Stage, approximately 6 million years in age (Bradley and
Landini 1984). However, these occurrences say little about the evolutionary biology
of hippoboscids (Dittmar et al. 2006). An alternative method is a molecular phylogenetic approach (Dittmar et al. 2006) that places the Hippoboscidae as the sistergroup to the Glossinidae, with an origin approximately at the same time as the
separation of the Nycteribiidae and Streblidae during the late Eocene (De Moya 2019).
Nycteribiidae (bat flies) are 12 genera of dorsoventrally flattened, spider-like
flies lacking eyes and wings that are obligate, host specific parasites on bats.
Nycteribiids have been implicated in the transmission of bat malaria in Africa
(Obame-Nkoghe et al. 2016). They lack a fossil record. Related to Nycteribiidae are
Streblidae (bat flies), consisting of 5 subfamilies and approximately 33 genera of
blood-feeding flies that are ectoparasitic specialists on a variety of bats (Lehane
1991). Some genera are involved in transmission of bat malaria (Obame-Nkoghe
et al. 2016). Although the oldest streblid is early Miocene in age (Poinar Jr and
Brown 2012), the clade is thought to have originated during the Eocene, based on
the fossil record of the closely related Glossinidae (Dittmar et al. 2006). From the
same early Miocene deposit, an unaffiliated streblid harbored Vetufebrus ovatus
(Haemospororida: Plasmodiidae), a bat malarial parasite (Poinar Jr 2011) that
apparently is not vectored by any known extant streblid (Leung 2017). This suggests that this particular parasitic relationship of malaria may be a disease that has
become extinct for streblid vectors and their bat hosts (compare van Dijk and De
Baets 2021 for other putative examples).
Oestroidea contains four families, Oestridae (bot flies), Calliphoridae (blow flies,
carrion flies, greenbottles), Mesembrinellidae (blow flies) and Mystacinobiidae
(New Zealand bat fly), which contain significant numbers of parasitic, bloodfeeding species. Parasitism likely originated five times in the Oestroidea, based on
genus-level phylogenies (Pape 2001; Kutty et al. 2001). As cyclorrhaphan calyptrates, Oestroidea are a recent clade that originated during the late Paleocene to
early Eocene epochs of the Cenozoic, about 60–40 million years ago (Cerretti et al.
2017). The most iconic of the oestroid parasitic families, Oestridae consist of about
160 species (Wood 2006) and contain the most conspicuous, diverse and noxious of
parasites, as they are internal parasites of mammals and frequently inhabit their
host’s flesh, occasionally using an intermediate vector such as a dipteran to complete their life cycle (Colwell et al. 2006). A typical life cycle is illustrated by the
reindeer warble fly, Hypoderma tarandi, documented on reindeer, Rangifer tarandus. The fly initially lays eggs during Arctic summer at the base of thin hairs adjacent the skin, followed by hatching and then penetration of the first-instar larva into
the skin, soon resulting in development of nodules (warbles) under the skin while
the host’s proteins from subcutaneous tissue are ingested by the larva (Asbakk et al.
2014). During the following Arctic winter, the larva continues to consume proteins,
C. C. Labandeira and L. Li