Palaeovespa florissantia.jpg
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''Palaeovespa'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
in the
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
subfamily
Vespinae The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets (the genus ''Vespa''), and the "yellowjackets" (genera ''Dolichovespula'' and ''Vespula''). The remaining genus, '' Provespa'', is a small, poorly ...
. The genus currently contains eight species, five from the Priabonian stage
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones an ...
in
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
, United States two from the
middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "da ...
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than ...
deposits of
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 subcontinent of Eurasia and it is located entirel ...
. and one species from the late Paleocene of France.


History and classification

The genus was first described by Dr. Theodore Cockerell in a 1906 paper published in the ''
Bulletin of the Museum of Comparative Zoology Bulletin or The Bulletin may refer to: Periodicals (newspapers, magazines, journals) * Bulletin (online newspaper), a Swedish online newspaper * ''The Bulletin'' (Australian periodical), an Australian magazine (1880–2008) ** Bulletin Debate ...
''. 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 ...
', meaning "old" and ''vespa'' from the genus ''
Vespa Vespa () is an Italian luxury brand of scooters and mopeds manufactured by Piaggio. The name means wasp in Italian. The Vespa has evolved from a single model motor scooter manufactured in 1946 by Piaggio & Co. S.p.A. of Pontedera, Italy to ...
'', the type genus of the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
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
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones an ...
. 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 Florissant. ''Palaeovespa'' gained another species, ''P. socialis'', in 2005 when George Poinar Jr. described a second species found in Baltic amber, while the eighth, and geologically oldest species, ''P. menatensis'', was published by Nel ''et al'' (2006).


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 Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella ...
''. Despite naming ''P. florissantia'' as the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
Cockerell noted that not all features of the genus were discernible in the ''P. forissantia''
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 sever ...
.


''P. florissantia''

At the time of description in 1906, 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 sever ...
for ''P. florissantia'' was the largest single hymenopteran specimen that Cockerell had seen from the Florissant Formation. He noted that holotype specimen ''
Museum of Comparative Zoology A museum ( ; plural museums or, rarely, musea) is a building or institution that cares for and displays a collection of artifacts and other objects of artistic, cultural, historical, or scientific importance. Many public museums make thes ...
(MCZ) number 2026'' was similar in appearance, on first examination, to a large
scoliid wasp The Scoliidae, the scoliid wasps, are a family of about 560 species found worldwide. They tend to be black, often marked with yellow or orange, and their wing tips are distinctively corrugated. Males are more slender and elongated than females, wi ...
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 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 . The
mesothorax The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
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 in recognition of his work on Colorado insects.


''P. baltica''

''P. baltica'' is the first of two ''Palaeovespa'' species described from
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than ...
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 The University of Königsberg (german: Albertus-Universität Königsberg) was the university of Königsberg in East Prussia. It was founded in 1544 as the world's second Protestant academy (after the University of Marburg) by Duke Albert of Pruss ...
amber collections. ''P. baltica'' is about in total length with forewings about . 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 shales on the Wilson Ranch near
Florissant, Colorado Florissant is an unincorporated town, a post office, and a census-designated place (CDP) located in and governed by Teller County, Colorado, United States. The CDP is a part of the Colorado Springs, CO Metropolitan Statistical Area. The Floriss ...
. The size and overall characters are similar to ''P. gillettei'' with an overall length of about and a forewing length of , 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 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 sever ...
is while the forewings are 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 long female specimen, number H-10-175 in the Poinar Amber Collection housed at
Oregon State University Oregon State University (OSU) is a public land-grant, research university in Corvallis, Oregon. OSU offers more than 200 undergraduate-degree programs along with a variety of graduate and doctoral degrees. It has the 10th largest engineering c ...
. 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 The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets (the genus ''Vespa''), and the "yellowjackets" (genera ''Dolichovespula'' and ''Vespula''). The remaining genus, '' Provespa'', is a small, poorly ...
species, the triangular arrangement of the
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
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
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
s, 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''.


''P. menatensis''

Oldest known species, described from the
Thanetian The Thanetian is, in the ICS Geologic timescale, the latest age or uppermost stratigraphic stage of the Paleocene Epoch or Series. It spans the time between . The Thanetian is preceded by the Selandian Age and followed by the Ypresian Age ( ...
Menat Formation, a
diatomite Diatomaceous earth (), diatomite (), or kieselgur/kieselguhr is a naturally occurring, soft, siliceous sedimentary rock that can be crumbled into a fine white to off-white powder. It has a particle size ranging from more than 3 μm to le ...
crater lake deposit in France


References

{{Taxonbar, from=Q7126519 Baltic amber Eocene insects Prehistoric insects of North America Fossil taxa described in 1906 Prehistoric Hymenoptera genera Florissant Formation