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''Protopone'' 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 of ants in the formicid subfamily
Ponerinae Ponerinae is a subfamily of ants in the Poneromorph subfamilies group, with about 1,600 species in 47 extant genera, including ''Dinoponera gigantea'' - one of the world's largest species of ant. Mated workers have replaced the queen as the fun ...
described from fossils 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 ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
. There are seven described species placed into the genus, ''Protopone? dubia'', ''Protopone germanica'', ''Protopone magna'', ''Protopone oculata'', ''Protopone primigena'', ''Protopone sepulta'', and ''Protopone vetula''. ''Protopone'' is one several Lutetian Ponerinae genera.


History and classification

When first described, ''Protopone'' was known from a single fossil insect included in a transparent chunk of Sakhalin amber. When the fossil was described it was part of the amber collections housed in the
Paleontological Institute, Russian Academy of Sciences The Paleontological Institute, Russian Academy of Sciences (PIN; russian: Палеонтологический институт РАН) in Moscow is among the world's largest paleontological institutes. An affiliate of the Russian Academy of Scienc ...
. The amber was recovered from fossil bearing rocks in the Sakhalin region of Eastern Russia. At the time of description, the amber was estimated to date from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
. Further research has revised the amber, which originates from the Lower Due Formation, to be of Middle Eocene age. The amber fossil was first studied by paleoentomologist Gennady Dlussky of the Russian Academy of Sciences, with his 1988
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 ...
for the species was published in ''Paleontologicheskii Zhurnal''. In 2012 an additional six species were identified from compression-impression fossils 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 ''Protopone'' specimens were collected from layers of
Lutetian The Lutetian is, in the geologic timescale, a stage or age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it is sometimes referred to as the Midd ...
Messel Formation The Messel Formation is a geologic formation in Hesse, central Germany, dating back to the Eocene epoch (about 47 MaJens Lorenz Franzen (2005). "The implications of the numerical dating of the Messel fossil deposit (Eocene, Germany) for mamma ...
rock in the
Messel pit The Messel pit (german: Grube Messel) is a disused quarry near the village of Messel ( Landkreis Darmstadt-Dieburg, Hesse) about southeast of Frankfurt am Main, Germany. Bituminous shale was mined there. Because of its abundance of well-preserv ...
World Heritage Site A World Heritage Site is a landmark or area with legal protection by an international convention administered by the United Nations Educational, Scientific and Cultural Organization (UNESCO). World Heritage Sites are designated by UNESCO for h ...
. The Messel formation is composed of brown coals, oil shales, and
bituminous shale Oil shale is an organic-rich fine-grained sedimentary rock containing kerogen (a solid mixture of organic chemical compounds) from which liquid hydrocarbons can be produced. In addition to kerogen, general composition of oil shales constitute ...
, which preserved numerous insects, fish, birds, reptiles, and terrestrial mammals as a notable lagerstätten. The area is a preserved maar lake which initially formed approximately 47 million years ago as the result of volcanic explosions. 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 ...
specimens for the six Messel species were preserved in the Senckenberg Research Station Messel fossil collections. The fossils were first described by Gennady Dlussky and Sonja Wedmann in a paper on the poneromorph ants of Messel. In the type description, Dlussky and Wedmann named the species ''P. ? dubia'', ''P. germanica'', ''P. magna'', ''P. oculata'', ''P. sepulta'', and ''P. vetula. ''Protopone'' is one of five extinct genera from three subfamilies which have species described from Messel Formation fossils by Dlussky and Wedmann in 2012. Three of the other genera ''
Cephalopone ''Cephalopone'' is an extinct genus of ants in the formicid subfamily Ponerinae described from fossils found in Europe. There are two described species placed into the genus, ''Cephalopone grandis'' and ''Cephalopone potens''. ''Cephalopone'' ...
'', '' Cyrtopone'', and '' Messelepone'' are also placed in Ponerinae. The genus ''
Casaleia ''Casaleia'' is an extinct genus of ants in the formicid subfamily Amblyoponinae described by Pagliano & Scaramozzino in 1990 from fossils found in Europe. The genus contains four species dating from the Eocene to Miocene, ''Casaleia eocenica'' ...
'' is placed in
Amblyoponinae Amblyoponinae is a subfamily of ants in the poneromorph subfamilies group containing 13 extant genera and one extinct genus. The ants in this subfamily are mostly specialized subterranean predators. Adult workers pierce the integument (non letha ...
, while the last genus ''
Pseudectatomma ''Pseudectatomma'' is an extinct genus of ants in the formicid subfamily Ectatomminae described by from fossils found in Europe. The genus contains two species dating from the Eocene, ''Pseudectatomma eocenica'' and ''Pseudectatomma striatula'' ...
'' is in Ectatomminae.


Description

The ''Protopone''
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
from Messel have approximate body lengths ranging between . Species have heads that are up to, but no shorter than the length of 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 ...
. The
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
is over 30% of the front area on the head, while the subtriangular mandibles are no more than 70% of the length of the head. The antennae sockets are widely placed on the head, in contrast to those seen in ''
Pachycondyla ''Pachycondyla'' is a ponerine genus of ants found in the Neotropics. Distribution ''Pachycondyla'' is currently distributed from southern United States to northern Argentina, but some fossil species (e.g. '' P. eocenica'' and '' P. lutzi'') a ...
'' which are close together. On both the hind and middle tibiae there are a pair of spurs, one simple and one pectinate. The claws of the pretarsals are simple. The petiole is rounded with a helcium that projects of the front face of the gasteral segment.


''P.? dubia''

''P.? dubia'' is known from a single partial fossil, consisting of a head, partial
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 ...
and fore-wing fragments. As such, the species name is based on the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
"dubia" meaning doubtful. The head looks to be narrower than it is long and has rounded corners and convex back edge. The eyes are placed just to the front of the heads midline, and the clypeus has a rounded front edge. The gena plates, below the eyes, are nearly as long as the eyes are wide. The subtriangular mandibles have many small teeth, over ten on each side. The widely placed antennae sockets and mandibles that are about 65% of the head length preclude placement into ''
Pachycondyla ''Pachycondyla'' is a ponerine genus of ants found in the Neotropics. Distribution ''Pachycondyla'' is currently distributed from southern United States to northern Argentina, but some fossil species (e.g. '' P. eocenica'' and '' P. lutzi'') a ...
'', ''Cephalopone'', ''Cyrtopone'', and ''Messelpone''. While the mandibles are similar in length to ''Pseudectatomma'', they are more gracile and the antenna scape is shorter. As such, the species was placed provisionally into ''Protopone''.


''P. germanica''

The species epithet was chosen by authors in recognition of Germany, the type locality. Two specimens were used to describe ''P. germanica'', a complete winged queen and a partial specimen of just the head. The body length of the holotype is with a node shaped petiole and a
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 ...
that is 1.3 times longer then the head. The eyes are placed just back from the midpoint of the head. The clypeus is rounded on it front edge and the
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
is about 45% as wide as the head. The antennae have a scape that reaches just past the back edge of the head, and there is an elongated segment at the antenna tip rather than a distinct club.


''P. magna''

Derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
"magnus", the species name ''magna'' was selected as a reference to the large size of the species. The two described gynes are preserved in lateral view, with the holotype being mostly complete and paratype specimen missing the head. The estimated body length for the holotype is with a long mesosoma. The head is covered with a distinct sculpturing of small pits, and the front edge of the clypeus is rounded as in other ''Protopone'' species. The antennae lack a club composed of the tip antenna segments, and the middle segments are nearly twice as long as they are thick. The scutum, on the front of the mesonotum, is shorter than seen in any of the other species. The petiole is just slightly higher than it is long and the front and top faces are rounded. The rear face is distinctly flattened.


''P. oculata''

The notably large eyes of ''P. oculata'' are the inspiration for the species name ''oculata'' which comes from the Latin "oculatus" meaning ''eyed''. The species is mid sized for the genus, with the holotype having a body length of and a mesosoma of . The ovoid eyes are large and placed slightly to the front of the heads mid-length. The mandibles are sub-triangular in outline with over 10 teeth on each side and are approximately 58% of the head length. The petiole node is a bit shorter than it is high and has rounded faces on each side.


''P. primigena''

The only specimen of ''P. primigena'' differs from the other species in several ways. The scape is shorter, not extending past the rear margin of the head capsule and the eyes are located closer to the mandibles than in the other species. The estimated body length is approximately with a petiole node that has a triangular outline.


''P. sepulta''

The mid sized species is known from a single fossil, with a body length of , making it the second largest of the species, only being smaller than ''P. magna''. The species name is from the Latin "sepultus" meaning buried, but the authors did not elaborate on the reason for the choice. While wings are preserved with the gyne, there is little detail visible of the wing venation. The head capsule is flat along the rear margin and the corners of the rear margin are rounded. The gena are slightly longer than the eyes are wide. The eyes are placed on or just behind the midpoint of the head. The antennae have a scape that is a little longer than that of ''P. primigena'', extending just past the back of the head.


''P. vetula''

The species name ''vetula'' is from the Latin "vetulus" meaning old or wizened. With a body length of it is the second smallest species, after ''P. primigena''. The head is wider than it is long and has eyes placed to the front of the midpoint. The mandible length is less than the head length. The mesosoma is long, 1.5 times longer than it is high.


References


External links

* * {{Taxonbar, from=Q18103620 †Protopone Fossil taxa described in 2012 Insects of Europe Fossil ant genera Prehistoric life of Europe Eocene insects Sakhalin amber