''Protohabropoda'' 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 ...

bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...

in the

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...

known from a fossil found in

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

. The genus currently contains a single described species ''Protohabropoda pauli''.

History and classification

''P. pauli'' was described from a solitary fossil, which is a compression-impression fossil pair preserved in layers of soft

sedimentary Sedimentary rocks are types of rock (geology), rock that are formed by the accumulation or deposition of mineral or organic matter, organic particles at Earth#Surface, Earth's surface, followed by cementation (geology), cementation. Sedimentati ...

rock. Along with other well preserved insect fossils, the ''P. pauli'' specimen was collected from layers of

Late Oligocene The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...

lacustrine A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much larger ...

rock belonging to the "calcaire de Campagne Calavon" sediments. The material is exposed along the northern slopes of the Luberon mountains near

Céreste Céreste (; Occitan: ''Ceirèsta'') is a commune in the Alpes-de-Haute-Provence department in southeastern France. It is known for its rich fossil beds in fine layers of "Calcaire de Campagne Calavon" limestone, which are now protected by the Pa ...

in Southern France. The sediments are reported as from a shallow paleolake that was formerly considered about 30 million years old and

Rupelian The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...

in age. Recent restudy of formation suggests the older Late Oligocene age. The paleoflora preserved in the shales suggest the lake was surrounded by a mixed-mesophytic forest though the vertebrate fauna found in the formation is more typical of a semi-arid environment. Specimens from the

Apoidea The superfamily (zoology), superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecidae, sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...

families are rather rare and not diverse, with '' Apis'' specimens being the most common. At the time of study, the holotype counterpart and part were part of the paleoentomology collections housed by the

Museum National d’Histoire Naturelle The French National Museum of Natural History, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is loca ...

. It was first studied by an international team of researchers headed by Manuel Dehon of the

University of Mons The University of Mons (french: Université de Mons) is a Belgian university located in the city of Mons, founded in 2009 by merging the Engineering Faculty of Mons (FPMs) and the University of Mons-Hainaut. The merging of the institutions wa ...

Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to th ...

, and the team's 2014

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 species was published in the

natural sciences Natural science is one of the branches of science concerned with the description, understanding and prediction of natural phenomena, based on empirical evidence from observation and experimentation. Mechanisms such as peer review and repeatab ...

journal '' PLOS ONE''. The genus name is 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 ...

word "protos" meaning ''first'' and the genus ''

Habropoda ''Habropoda'' is a genus of anthophorine bees in the family Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumb ...

'' of which ''Protohabropoda'' is most similar. 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 ...

''pauli'' is a patronym coined in honor of Paul Léon Victor Vigot, a young bee systematics enthusiast. The size and shape of the

pterostigma The pterostigma (plural: pterostigmata) is a group of specialized cells in the outer insect wing, wings of insects, which are often thickened or coloured, and thus stand out from other cells. It is particularly noticeable in dragonfly, dragonfli ...

are similar to those of the Apinae tribes

Anthophorini The Anthophorini are a large tribe in the subfamily Apinae of the family Apidae. Species in this tribe are often referred to as digger bees, although this common name is sometimes also applied to members of the tribe Centridini. It contains over ...

and

Centridini The Centridini are a tribe of large apid bees, many of which possess adaptations for carrying floral oils rather than (or in addition to) pollen or nectar. The floral oils are often gathered from plants of the family Malpighiaceae, though other ...

though the rounded shape of the wing and the sizes of the cells formed by the wing veins make it closer to Anthophorini. In the tribe, ''P. pauli'' is identified from the living genera ''

Anthophora The bee genus ''Anthophora'' is one of the largest in the family Apidae, with over 450 species worldwide in 14 different subgenera. They are most abundant and diverse in the Holarctic and African biogeographic regions. All species are solitary, t ...

'' and ''

Amegilla ''Amegilla'' is a large genus of bees in the tribe Anthophorini. Several species have blue metallic bands on the abdomen, and are referred to as "blue-banded bees" and "digger bees". One of their names in english, "digger bees" is given to them ...

'' by the positioning of veins and cells. and the genus ''Habropoda'' is distinguished by vein length. ''P. pauli'' is one of four bee species described by Dehon and team in the ''PLOS ONE'' article, the others being ''

Andrena antoinei ''Andrena antoinei'' is an extinct species of mining bee in the family Andrenidae described from a single fossil found in a Late Oligocene lake in present-day France that existed in semi-arid conditions. History and classification ''A. antoinei ...

'', ''

Bombus cerdanyensis ''Bombus cerdanyensis'' is an extinct species of bumble bee in the family Apidae known from a fossil found in Europe. History and classification ''Bombus cerdanyensis'' was described from a solitary fossil, which is a compression-impression fos ...

'', and '' Euglossopteryx biesmeijeri''.

Description

The ''P. pauli'' fossil is a female preserved with a dorsal view of the body, head twisted upward showing the face, the right forewing outstretched, and hindwings missing. The overall body length is not determinable due to positioning of the body and the twisting of the head although the

mesosoma The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. In hymenopterans of t ...

is . The incompletely preserved antennae are not distinct enough to determine

flagellomere Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for sensing in arthropods. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in form but are always made of one o ...

numbers or lengths. The

metasoma The metasoma is the posterior part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory system, and circula ...

is long in side view, and the two sections that are distinctly preserved show a dense coating of hairs. The legs are not well preserved, being partly covered by the body segments though the areas that are visible show a dense coating of setae. The long forewings have a one marginal cell and three cells below that called the submarginal cells. The marginal cell has very little tapering near the apical end, which is distinctly rounded. The second and third submarginal cells together are longer than the first submarginal cell, which is the longest of the three cells. The

is present, with parallel sides and, like the marginal cell, no narrowing of the width.

References

External links

* * {{Taxonbar, from=Q21808757 † Fossil taxa described in 2014 Prehistoric insects of North America Eocene insects Fossil bee genera Taxa named by Michael S. Engel