''Enischnomyia'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

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 ho ...

. 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 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 means "less recen ...

fossil found on

Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti or Quisqueya) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and th ...

. ''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 incl ...

. The amber is fossil resin that was produced by the extinct ''

Hymenaea protera ''Hymenaea protera'' is an extinct prehistoric leguminous tree, the probable ancestor of present-day ''Hymenaea'' species. Most neotropical ambers come from its fossilized resin, including the famous Dominican amber. ''H. protera'' once grew in ...

'', which formerly grew on

, across northern South America and up to southern Mexico. The amber dates from the

Burdigalian The Burdigalian is, in the geologic timescale, an age (geology), age or stage (stratigraphy), stage in the early Miocene. It spans the time between 20.43 ± 0.05 annum, Ma and 15.97 ± 0.05 Ma (million years ago). Preceded by the Aquitanian (sta ...

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 The La Toca Formation is a geologic formation in the northern and eastern part of the Dominican Republic. The formation, predominantly an alternating sequence of marls and turbiditic sandstones, breccias and conglomerates, is renowned for the p ...

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 island in Haiti, and to the southeast through lowlands to where it rises as the Sierra d ...

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 wit ...

. 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 several ...

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 co ...

, Corvallis. The holotype fossil was first studied by entomologist

George Poinar Jr. George O. Poinar Jr. (born April 25, 1936) is an American entomologist and writer. He is known for popularizing the idea of extracting DNA from insects fossilized in amber, an idea which received widespread attention when adapted by Michael Cr ...

of Oregon State University, and Alex Brown of Berkeley, California, with their 2012

type description A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have be ...

of the new genus and species being published in the journal ''

Systematic Parasitology ''Systematic Parasitology'' is a monthly peer-reviewed medical journal covering all aspects of the taxonomy and systematics of parasites. It was established in 1979 and is published by Springer Science+Business Media. The editor-in-chief is Anet ...

''. 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 In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

''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 cer ...

''. In 2005 additional bat hairs were reported in the amber specimen entombing the extinct kissing-bug ''

Triatoma dominicana ''Triatoma dominicana'' is an extinct species of assassin bug in the subfamily Triatominae, the kissing bugs known from early Miocene Burdigalian stage Dominican amber deposits on the island of Hispaniola. The species is known from a single 5th ...

'', itself a host to the extinct ''

Trypanosoma antiquus ''Trypanosoma antiquus'' is an extinct species of kinetoplastid (class Kinetoplastida), a monophyletic group of unicellular parasitic flagellate protozoa. The genus name is derived from the Greek ''trypano'' (borer) and ''soma'' (body) beca ...

''. 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

oocyst 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 i ...

s 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 i ...

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. Th ...

, 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