''Protohabropoda'' 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 ...

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 ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

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 continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. 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 formed by the cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or deposited at Earth's surface. Sedime ...

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 geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the pro ...

lacustrine A lake is often a naturally occurring, relatively large and fixed body of water on or near the Earth's surface. It is localized in a basin or interconnected basins surrounded by dry land. Lakes lie completely on land and are separate from t ...

rock belonging to the "calcaire de Campagne Calavon" sediments. The material is exposed along the northern slopes of the Luberon mountains near Céreste 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 age (geology), ages or the lower of two stage (stratigraphy), stages of the Oligocene epoch (geology), Epoch/series (stratigraphy), Series. It spans the time between . It is preceded b ...

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 Apoidea is a major group (of over 30 000 species) within the Hymenoptera, which includes two traditionally recognized lineages, the "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. It was first studied by an international team of researchers headed by Manuel Dehon of the

University of Mons The University of 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 was achieved following a g ...

Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...

, and the team's 2014

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 species was published in the

natural sciences Natural science or empirical 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 ...

journal '' PLOS ONE''. The genus name is a combination of the

Greek Greek may refer to: 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 of all kno ...

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 bumbl ...

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

''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 wings of insects, which are often thickened or coloured, and thus stand out from other cells. It is particularly noticeable in dragonflies, but present also in ...

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 othe ...

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 Africa Africa is the world's second-largest and sec ...

'' and ''

Amegilla ''Amegilla'' is a large genus of bees in the tribe Anthophorini. It has been split into many subgenera, including '' Asaropoda'', ''Micramegilla,'' '' Notomegilla'' and '' Zonamegilla''. Description ''Amegilla'' are generally medium-sized to ...

'' 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'', 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. Wasps, bees and a ...

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

flagellomere An antenna (plural: antennae) is one of a pair of appendages used for sensing in arthropods. Antennae are sometimes referred to as ''feelers''. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in ...

numbers or lengths. The

metasoma The metasoma is the posterior part of the body, or tagma (biology), 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 sy ...

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

† A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...

Fossil taxa described in 2014 Prehistoric insects of North America Eocene insects Fossil bee genera Taxa named by Michael S. Engel Monotypic prehistoric insect genera