''Vetufebrus'' is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of haemospororida in the family

Plasmodiidae The Plasmodiidae are a family of apicomplexan parasites, including the type genus ''Plasmodium'', which is responsible for malaria. This family was erected in 1903 by Mesnil and is one of the four families in the order Haemospororida. Diagnos ...

. At the time of its description the new genus comprised a single species ''Vetufebrus ovatus'' 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 ...

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 in ...

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

found on

Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...

. ''V. ovatus'' was vectored by '' Enischnomyia stegosoma'', the first fossil streblid bat fly described from a fossil, and the only member of the subfamily Nycterophiliinae described from Hispaniola. ''V. ovatus'' is the first instance of a Streblidae bat fly as a host for a

malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...

l parasite.

History and classification

''Vetufebrus ovatus'' was described based on a group of fossilized specimens which are preserved as in the host batfly, itself an

inclusion Inclusion or Include may refer to: Sociology * Social inclusion, action taken to support people of different backgrounds sharing life together. ** Inclusion (disability rights), promotion of people with disabilities sharing various aspects of lif ...

in a transparent chunk of

. The amber is fossil resin that was produced by the extinct '' Hymenaea protera'', 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 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 The Yanigua Formation is a Formation (geology), geologic formation in Dominican Republic. The lagoonal claystones and marls preserve fossils dating back to the Miocene Period (geology), period. The amber specimen was collected from the LaBúcara amber mine in the

Dominican Republic The Dominican Republic is a country located on the island of Hispaniola in the Greater Antilles of the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean. It shares a Maritime boundary, maritime border with Puerto Rico to the east and ...

. At the time of description, the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

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 university, public Land-grant university, land-grant research university in Corvallis, Oregon, United States. OSU offers more than 200 undergraduate degree programs and a variety of graduate and doctor ...

Corvallis, Oregon Corvallis ( ) is a city in and the county seat of Benton County, Oregon, Benton County in central western Oregon, United States. It is the principal city of the Corvallis, Oregon Metropolitan Statistical Area, which encompasses all of Benton Co ...

. 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 Crich ...

of Oregon State University with his 2011

type description A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...

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

Parasites & Vectors ''Parasites & Vectors'' is a peer-reviewed open-access medical journal published by BioMed Central. The journal publishes articles on the biology of parasites, parasitic diseases, intermediate hosts, vectors and vector-borne pathogens. ''Parasites ...

''. The genus name, ''Vetufebrus'' was derived from a combination of the

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

words ''vetus'' meaning "old" and ''febris'' meaning "fever". The

specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

''ovatus'' was coined from the Latin word ''ovatus'' meaning "ovate", an allusion to the shape of the oocysts.

Paleobiology and parasite vectoring

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. Together with their close relatives the Streblidae, they are known as "bat flies". As the latter do not seem to be a monophyletic group, it is conceivable that bat flies ca ...

, with no recorded association between living malaria plasmodiids and streblid bat flies. Of the four genera of bat malaria ''

Hepatocystis ''Hepatocystis'' is a genus of parasites transmitted by midges of the genus ''Culicoides''. Hosts include Old World primates, bats, hippopotamus and squirrels. This genus is not found in the New World. The genus was erected by Levaditi and Schoen, ...

'', ''

Nycteria ''Nycteria'' is a genus of protozoan parasites that belong to the phylum Apicomplexa. It is composed of vector-borne haemosporidian parasites that infect a wide range of mammals such as primates, rodents and bats. Its vertebrate hosts are bats. ...

'', ''

Plasmodium ''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a Hematophagy, blood-feeding insect host (biology), host which then inj ...

'', and '' Polychromophilus'', the first three are confined to

old world The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...

bats, and only ''Polychromophilus'' is present in the

new world The term "New World" is used to describe the majority of lands of Earth's Western Hemisphere, particularly the Americas, and sometimes Oceania."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: ...

. ''Vetufebrus ovatus''

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 ...

s are preserved in the mid gut of '' Enischnomyia stegosoma'' 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 organis ...

s are present in the oocysts along with in the salivary ducts. The sporozoites preserved in the salivary glands and ducts are similar in size to those in the oocysts, which indicate that ''V. ovatus'' was successfully vectored to the flies host. The infection association was the first instance of a

Streblidae The Streblidae are a Family (biology), family of fly, flies in the Taxonomic rank, superfamily Hippoboscoidea, and together with their relatives the Nycteribiidae, are known as bat flies. They are winged or wingless ectoparasites of bats, and oft ...

family bat fly acting as host and vector for a malarial parasite. The oocysts of ''V. ovatus'' are smaller and ovoid in shape while mature ''Polychromophilus'' species oocysts are rounded and larger overall. Poinar notes that the oocysts in ''V. ovatus'' might have been immature, and the sporozoites in the salivary ducts could have been left from a prior infection. The overall sporozoite morphology is similar to those of ''Polychromophilus'' species, suggesting the possibility of ''Vetufebrus'' being an early lineage for ''Polychromophilus'', though the oocysts are not consistent. The amber entombing ''V. ovatus'' and ''E. stegosoma'' contains no preserved evidence of what its host animal may have been. However member species of Streblidae are bat parasites, with Nycterophiliinae subfamily 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 ha ...

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 ...

'', its-self 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) because ...

''.

Description

Both the oocysts are a brown in coloration and are between long by wide. Both oocysts have a thin surrounding membrane, multiple dark colored cells with nuclei and a number of developing sporozoites. The sporozoites in the oocysts range between , while the sporocytes in the salivary glands are slightly larger, and have a rounded stubby outline.

References

{{Taxonbar, from=Q7923842 Apicomplexa genera Haemosporida Burdigalian life Neogene Dominican Republic Fossils of the Dominican Republic Dominican amber Fossil taxa described in 2011 Taxa named by George Poinar Jr.