Enischnomyia With Vetufebrus 1
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''Enischnomyia'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of bat fly in the family
Streblidae The Streblidae are a family of flies in the superfamily Hippoboscoidea, and together with their relatives the Nycteribiidae, are known as bat flies. They are winged or wingless ectoparasites of bats, and often have long legs. They appear to be ...
. At the time of its description the new genus comprised a single species, ''Enischnomyia stegosoma'', known from a single
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
fossil found on Hispaniola. ''E. stegosoma'' was the first fossil streblid bat fly described from a fossil, and the only member of the subfamily Nycterophiliinae described from Hispaniola. The species is host for the plasmodiid '' Vetufebrus ovatus'' preserved in its salivary glands and midgut.


History and classification

''Enischnomyia stegosoma'' was described based on a single fossilised specimen which is preserved as an
inclusion Inclusion or Include may refer to: Sociology * Social inclusion, aims to create an environment that supports equal opportunity for individuals and groups that form a society. ** Inclusion (disability rights), promotion of people with disabiliti ...
in a transparent chunk of
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil inc ...
. The amber is fossil resin that was produced by the extinct '' Hymenaea protera'', which formerly grew on Hispaniola, across northern South America and up to southern Mexico. The amber dates from the
Burdigalian The Burdigalian is, in the geologic timescale, an age or stage in the early Miocene. It spans the time between 20.43 ± 0.05 Ma and 15.97 ± 0.05 Ma (million years ago). Preceded by the Aquitanian, the Burdigalian was the first and longest w ...
stage (20.43 ± 0.05 to 15.97 ± 0.05 million years ago) of the Miocene, and is recovered from sections of the La Toca Formation in the
Cordillera Septentrional The Cordillera Septentrional is a mountain range that runs parallel to the north coast of the Dominican Republic, with extensions to the northwest as Tortuga (Haiti), Tortuga island in Haiti, and to the southeast through lowlands to where it rises ...
and the Yanigua Formation in the Cordillera Oriental. The specimen was collected from the LaBúcara amber mine in the
Dominican Republic The Dominican Republic ( ; es, República Dominicana, ) is a country located on the island of Hispaniola in the Greater Antilles archipelago of the Caribbean region. It occupies the eastern five-eighths of the island, which it shares with ...
. At the time of description, the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
specimen, number "No. D-7-239", was preserved in the Poinar Amber collections, housed at
Oregon State University Oregon State University (OSU) is a public land-grant, research university in Corvallis, Oregon. OSU offers more than 200 undergraduate-degree programs along with a variety of graduate and doctoral degrees. It has the 10th largest engineering c ...
, Corvallis. The holotype fossil was first studied by entomologist George Poinar Jr. of Oregon State University, and Alex Brown of Berkeley, California, with their 2012 type description of the new genus and species being published in the journal '' Systematic Parasitology''. The genus name, ''Enischnomyia'' was derived from a combination of the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words ''myia'' meaning "fly" and ''enischnos'' meaning "thin" or "slight". The specific epithet ''stegosoma'' is coined from the Greek words ''stenos'' and ''soma'' meaning "narrow" and "body" respectively.


Paleobiology and parasite vectoring

The amber entombing ''E. stegosoma'' contains no preserved evidence of what its host animal may have been. However member species of Streblidae are bat parasites, with the Nycterophiliinae species being obligate external parasites that feed on the blood of bats, so it is suggested ''E. stegosoma'' was the same. At least two bat fur fossils had been described from Dominican amber prior to the description of ''E. stegosoma''. A larval
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
brush-footed butterfly with a single hair stuck to a rear spine was described in 1998, with the hair noted to be similar to those of the bat genus ''
Eptesicus ''Eptesicus'' is a genus of bats, commonly called house bats or serotine bats, in the family Vespertilionidae. The genus name is likely derived from the Greek words ''ptetikos'' 'able to fly' or ''petomai'' 'house flier', although this is not ce ...
''. In 2005 additional bat hairs were reported in the amber specimen entombing the extinct kissing-bug '' Triatoma dominicana'', itself a host to the extinct '' Trypanosoma antiquus''. Based on the flattened and rather flea like body plus inflated front femurs, simplified wing structure and location of origin, the genus was placed into the bat fly subfamily Nycterophiliinae. The flattened body and front legs were likely used to burrow into the bat's fur to reach skin for feeding, as is done by modern '' Nycterophilia coxata''. Unlike the modern nycterophiliines, which lower their entire head to the skin to feed, ''E. stegosoma'' had an elongated ' labium which was most likely lowered to the skin instead. No members of the subfamily are native to Hispaniola, which is
depauperate A depauperate ecosystem is an ecosystem which is lacking in numbers or variety of species, often because it lacks enough stored chemical elements and resources required for life. Thus, depauperate ecosystems often cannot support rapid growth of f ...
of bat-flies, having two genera, '' Streba'' and '' Tricholobius'' and five total species. Preserved in the mid gut and salivary ducts of ''E. stegosoma'' are oocysts and
sporozoite Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism is ...
s of the plasmodiid '' Vetufebrus ovatus''. The association was the first instance of a streblid bat fly acting as host and vector for a malarial parasite. Extant bat malaria causing plasmodiids are transmitted by species of the bat fly family
Nycteribiidae Nycteribiidae is a family of the true fly superfamily Hippoboscoidea are known as "bat flies", together with their close relatives the Streblidae. As the latter do not seem to be a monophyletic group, it is conceivable that bat flies cannot be u ...
, with no recorded association between living malaria plasmodiids and streblid bat flies.


Description

The holotype male is with wings that are long by wide. The head on a prominent neck is partially covered by the expanded procoxae of the front most legs. There are no
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
and the compound eyes at the antennae bases are reduced in size to only three facets. The antennae have two very modified segments, a pyramidal basal scape that is short and attached to the head. At the tip of the scape are both the flagellum, and the pedicel with ringed tube-like lower section and enlarged apex. On the pedicel apex are seven spine-like
setae In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
, four small ones and three elongated ones. The flagellum is placed below the pedicel and forms a plumose arista. The labium has two small setae on the underside of the fleshy, small labella along with five more setae on the terminal edge. The labrum has a tube shape and the basal theca is slightly enlarged.


References

{{Taxonbar, from=Q21364263 Prehistoric Diptera Miocene insects Burdigalian life Prehistoric insects of the Caribbean Fossils of the Dominican Republic Dominican amber Fossil taxa described in 2012