Palaeovespa
Encyclopedia
Palaeovespa is an extinct genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of wasp in the Vespidae subfamily Vespinae
Vespinae
The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets , and the "yellowjackets" . The remaining genus, Provespa is a small, poorly-known group of nocturnal wasps from Southeast Asia. One genus, Palaeovespa, has been described from the Eocene fossil...

. The genus currently contains seven species, five from the Priabonian
Priabonian
The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene epoch or series. It spans the time between and...

 stage Florissant Formation in Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...

, USA and two from the middle Eocene Baltic amber
Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite, with about 80% of the world's known amber found there. It dates from 44 million years ago...

 deposits of Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

.

History and classification

The genus was first described by Dr. Theodore Cockerell
Theodore Dru Alison Cockerell
Theodore Dru Alison Cockerell was an American zoologist, born at Norwood, England, and brother of Sydney Cockerell. He was educated at the Middlesex Hospital Medical School, and then studied botany in the field in Colorado in 1887-90...

 in a 1906 paper published in the Bulletin of the Museum of Comparative Zoology. The genus name is a combination of the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 , meaning "old" and vespa from the genus Vespa
Hornet
Hornets are the largest eusocial wasps; some species can reach up to in length. The true hornets make up the genus Vespa and are distinguished from other vespines by the width of the vertex , which is proportionally larger in Vespa and by the anteriorly rounded gasters .- Life cycle :In...

, the type genus of the family Vespidae where Palaeovespa is placed.

Along with the genus description, the paper contained the description of the type species P. florissantia, P. scudderi and P. gillettei all from the Florrisant Formation. Cockerell described a fourth species, P. baltica in 1909 from a specimen in Baltic amber. Five years later, in 1914, Cockerell described another species P. wilsoni from Florissant. In 1923, P. relecta was named by Cockerell, bring the species count to six, with five described from from Florissant. Palaeovespa gained one more species, P. socialis, in 2005 when George Poinar, Jr.
George Poinar, Jr.
George O. Poinar, Jr. is an 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 Crichton for the book and movie Jurassic Park.Poinar earned a B.S. and M.S. at Cornell...

 described a second species found in Baltic amber.

Description

Palaeovespa is most similar to the extant genus Vespa, with which it shares many similar features such as a broad rounded thorax with a sessile abdomen that is broad at the base. The genus however possesses wing venation that is closer to the more primitive genus Polistes
Polistes
Wasps of the cosmopolitan genus Polistes are the most familiar of the polistine wasps, and are the most common type of paper wasp. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies...

. Despite naming P. florissantia as the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 Cockerell noted that not all features of the genus were discernible in the P. forissantia holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

.

P. florissantia

At the time of description in 1906, the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 for P. florissantia was the largest single hymenopteran
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

 specimen that Carpenter had seen from the Florissant Formation. He noted that holotype specimen Museum of Comparative Zoology
Museum of Comparative Zoology
The Museum of Comparative Zoology, full name "The Louis Agassiz Museum of Comparative Zoology", often abbreviated simply to "MCZ", is a zoology museum located on the grounds of Harvard University in Cambridge, Massachusetts. It is one of three museums which collectively comprise the Harvard Museum...

 (MCZ) number 2026
was similar in appearance, on first examination, to a large scoliid wasp
Scoliidae
Scoliidae, the scoliid wasps, is a small family represented by 6 genera and about 20 species in North America, but they occur worldwide, with a total of around 300 species. They tend to be black, often marked with yellow or orange, and their wing tips are distinctively corrugated...

 but that the specimen was undoubtedly a vespid. P. florissantia was apparently broadly black in color with the hind margins of the abdominal segments pallid in coloration, but displaying no distinct markings. The wings were possibly reddish in coloration. Though the wings are folded, the very long first discoidal, which is unique to members of Vespidae, is visible. The specific epithet is a reference to the Florissant Formation, which produced the specimen.

P. scudderi

P. scudderi, unlike P. florissantia, was described from two specimens. However only holotype specimen MCZ number 2027 was noted as a type specimen. The second specimen, MCZ number 7738, while used in the description, was not noted as a paratype. The species has eyes that are deeply emarginate like modern Vespa species, but unlike the modern species the eye segment above the emargination is very close to the size of the segment below. Also notable to P. scudderi are the strong vertical striations on the lateral prothorax lobes, a feature not seen in the other Palaeovespa species that Cockerell described in 1906. The head and thorax were dark, most likely black, in life, and the middle leg was black down to the apical third of the femur, at which point it shifts to a lighter tone, probably yellow or red. Cockerell named P. scudderi for Samuel H. Scudder
Samuel Hubbard Scudder
Samuel Hubbard Scudder was an American entomologist and palaeontologist.Born in Boston, Massachusetts, Scudder may be most widely known for his essay on the importance of first-hand, careful observation in the natural sciences...

 who was the first North American paleoentomologist and collector of numerous Florissant Formation insects.

P. gillettei

Of the three species first described in 1906, P. gillettei was described from the most specimens, with five specimens being referred to the species in addition to the holotype specimen, MCZ number 2028. This species is smaller and more delicate then either P. florissantia or P. scudderi with a body length of approximately 14.5 millimetre (0.570866141732284 in). The mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...

 possibly displayed two longitudinal stripes of yellow. The vein structure of the wings is very similar to that of P. scudderi, while the base of the abdomen is smaller than that of modern Vespa species. There is little flaring in the base of the abdomen, with the broadest part being just past the middle. The costal cell of the forewing shows a distinct dark area similar to the dark area on modern Vespa wings. The species was named in honor of Clarence P. Gillette
Clarence Preston Gillette
Clarence Preston Gillette was an American entomologist....

 in recognition of his work on Colorado insects.

P. baltica

P. baltica is the first of two Palaeovespa species described from Baltic amber
Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite, with about 80% of the world's known amber found there. It dates from 44 million years ago...

 and is known only from the type specimen, a single adult female. The species was named in 1909 by Cockerell in a paper describing a number of fossil Hymenoptera housed in the University of Königsberg
University of Königsberg
The University of Königsberg was the university of Königsberg in East Prussia. It was founded in 1544 as second Protestant academy by Duke Albert of Prussia, and was commonly known as the Albertina....

 amber collections. P. baltica is about 16 millimetre (0.62992125984252 in) in total length with forewings about 11 mm (0.433070866141732 in). The abdomen is "rusty red" in color with the underside darkening towards the apex. The legs are a dark "rusty red" or "yellowish rusty red", while the wings are "soot brown" in color. In the description of P. socialis, Poinar noted the inclusion of the two Baltic amber species in Palaeovespa to be dubious.

P. wilsoni

In 1914, five years after the publication of P. baltica and eight years after the description of the genus, Cockerell published the fourth Florissant formation species P. wilsoni. This species was described from a single female specimen recovered from shale
Shale
Shale is a fine-grained, clastic sedimentary rock composed of mud that is a mix of flakes of clay minerals and tiny fragments of other minerals, especially quartz and calcite. The ratio of clay to other minerals is variable. Shale is characterized by breaks along thin laminae or parallel layering...

s on the Wilson Ranch near Florissant, Colorado
Florissant, Colorado
Florissant is a census-designated place and a U.S. Post Office in Teller County, Colorado, United States. The population as of the 2010 Census was 104.Florissant, Colorado, was named after Florissant, Missouri, the hometown of the founding family...

. The size and overall characters are similar to P. gillettei with an overall length of about 15 millimetre (0.590551181102362 in) and a forewing length of 9 mm (0.354330708661417 in), the head and thorax are black while the abdomen and legs are pale, possibly having been yellow in life. The wings are glassy transparent ("hyaline"), with nearly colorless veins, in contrast to the wings of P. gillettei which have a dark area in the costal region. The wing vein characters were considered distinct enough to discount the possibly of the P. wilsoni holotype being a variant specimen of P. gillettei. Cockerell coined the specific name "wilsoni" in honor of the owners of the Wilson ranch who aided in the fossil-collecting efforts.

P. relecta

In 1923 Cockerell published his final species of Palaeovespa to be described from the Florissant Formation, P. relecta. The type and only known specimen is part of the Colorado Museum of Natural History collections and was loaned to Cockerell by the museum's director at that time, Jesse D. Figgins. While Cockerell considered the species to be close to P. gillettei, P. relecta is larger. The total length of the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 is 17.5 millimetre (0.688976377952756 in) while the forewings are 10.5 mm (0.413385826771654 in) long. P. relecta is noted for having abdominal coloration close to that of modern Vespa, possessing an abdomen that is mostly black with narrow light sutural bands. The head and thorax are black, the antenna ferruginous and darkening at the extreme base. The wings are hyaline with a dusky tint along the upper margin and lightly ferruginous veins.

P. socialis

P. socialis is the newest addition to the genus and the second species to be described from a Baltic amber specimen. The holotype is a 12.8 millimetre (0.503937007874016 in) long female specimen, number H-10-175 in the Poinar Amber Collection housed at Oregon State University
Oregon State University
Oregon State University is a coeducational, public research university located in Corvallis, Oregon, United States. The university offers undergraduate, graduate and doctoral degrees and a multitude of research opportunities. There are more than 200 academic degree programs offered through the...

. The head is light in coloration with yellow bands around the eyes and mouth parts. The mesosoma is mostly black while the tergites of the abdomen are black on the front portion and yellow on the back portion. The specific name "socialis" is in reference to the probable social habits and nature of the species. Of the two other described Baltic amber Vespinae
Vespinae
The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets , and the "yellowjackets" . The remaining genus, Provespa is a small, poorly-known group of nocturnal wasps from Southeast Asia. One genus, Palaeovespa, has been described from the Eocene fossil...

 species, the triangular arrangement of the ocelli separate P. socialis from Vespa dasypodia. While the size difference between P. baltica and P. socialis could be due to the specimen's being from different castes of the same species; the coloration and wing vein differences support the separation into two species. The holotype specimen is preserved with a small section of tissue with prolegs with crochets clenched in the mandibles, indicating the larvae used caterpillars as a source of protein. In his description of the species George Poinar notes the possibility that the Baltic amber species may not belong in Palaeovespa.
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