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''Parastylotermes'' 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 ...
of
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
in the
Isoptera Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
family
Stylotermitidae Stylotermitidae is a family of termites in the order Blattodea Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence sugge ...
known from North America, Europe, and India. The genus contains five described species, ''Parastylotermes calico'', ''Parastylotermes frazieri'', ''Parastylotermes krishnai'', ''Parastylotermes robustus'', and ''Parastylotermes washingtonensis''.


History and classification

''Parastylotermes'' was first described by
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
s Thomas E. Snyder and
Alfred E. Emerson Alfred Edwards Emerson, Jr. (December 31, 1896 – October 3, 1976) was an American biologist, Professor of Zoology at the University of Chicago, a noted entomologist and leading authority on termites. Life and work Emerson was born in Ithaca ...
in the 1949 paper ''Catalog of the termites (Isoptera) of the world''. Prior to 1949, both ''P. washingtonensis'' and ''P. robustus'' were placed in separate modern species. Snyder and Emerson placed ''Parastylotermes'' in Stylotermitidae, which at that time was considered a subfamily, Stylotermitinae, of the family
Rhinotermitidae Rhinotermitidae is a family of termites (Isoptera). They feed on wood and can cause extensive damage to buildings or other wooden structures. About 345 species are recognized, among these are severe pests such as '' Coptotermes formosanus'', ''Co ...
.


Species


''P. robustus''

''P. robustus'' was the first species to be described, having been published by Baron Kurt von Rosen as ''Leucotermes robustus'' in 1913. Fossils of the species were recovered from the Middle Eocene,
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 ...
, "Blue Earth"
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 1 ...
deposits, with three
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the f ...
s and four isolated wings known. Of the five known specimens, the original type series of fossils that von Rosen used in describing the species were held in the Bayerische Staatssammlung für Paläontologie und Geologie collections and are thought destroyed during
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
. Von Rosen noted in his type description of the species that he was uncertain of its genus placement. Since then the species has been moved several first to ''Leucotermes (Reticulitermes) robustus'', then to ''Reticulitermes robustus'' when ''
Reticulitermes ''Reticulitermes'' is a termite genus in the family Rhinotermitidae. They are found in most temperate regions on Earth including much of Asia and the Middle East, Western Europe, and all of North America. Caste descriptions ''Reticulitermes'' ...
'' was changed from a subgenus to a full genus. Finally the species was transferred to ''Parastylotermes'' by Snyder and Emerson in 1949. ''P. robustus'' and ''P. krishnai'' are the only two species which are known from more than isolated wings. ''P. robustus'' can be distinguished from ''P. washingtonensis'' by is age and its shorter, wider, wings which are estimated to be .


''P. washingtonensis''

''P. washingtonensis'', described by Snyder in 1931 as ''Stylotermes washingtonensis'', is 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 specimen ...
for ''Parastylotermes''. The species, like ''P. frazieri'' and ''P. calico'', is known from a single isolated wing. The front right fore-wing was found in sediments of the
Latah Formation The Latah Formation is a series of late Miocene lacustrine sedimentary deposits which outcrop in eastern Washington and northwestern Idaho. The lake beds are interbedded with igneous rock of the Columbia River Basalt Group. The formation was ...
near
Spokane, Washington Spokane ( ) is the largest city and county seat of Spokane County, Washington, United States. It is in eastern Washington, along the Spokane River, adjacent to the Selkirk Mountains, and west of the Rocky Mountain foothills, south of the Canada ...
. The formation is dated to the Upper Miocene placing as similar in age to both other western North American species, both being middle Miocene in age. The specimen is housed as specimen ''MCZ 2943 ab'' in the
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 these ...
on the
Cambridge, Massachusetts Cambridge ( ) is a city in Middlesex County, Massachusetts, United States. As part of the Boston metropolitan area, the cities population of the 2020 U.S. census was 118,403, making it the fourth most populous city in the state, behind Boston, ...
grounds on
Harvard University Harvard University is a private Ivy League research university in Cambridge, Massachusetts. Founded in 1636 as Harvard College and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher le ...
. At an estimated length from the costal suture to the tip and a full , is one of the largest species of ''Parastylotermes'', with ''P. calico'' estimated to have been slightly larger. The structuring of the median and cubitus veins are cited as the major differences between ''P. washingtonensis'' and ''P. calico''.


''P. frazieri''

''P. frazieri'' was the third species of ''Parastylotermes'' to be described and is known from a single wing collected in 1954 by T. H. McCulloh of the
United States Geological Survey The United States Geological Survey (USGS), formerly simply known as the Geological Survey, is a scientific agency of the United States government. The scientists of the USGS study the landscape of the United States, its natural resources, ...
and deposited into the
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums and education and research centers, the largest such complex in the world, created by the U.S. government "for the increase and diffusion of knowledge". Founded ...
as specimen number 62383. The wing was recovered from an calcium carbonate nodule recovered from the Frazier borax mine in Ventura county California. When Thomas Snyder described the specimen in 1955 he was uncertain whether the wing was in fact a fore-wing. The wing is long and wide and differs from ''P. washingtonensis'' in the placement of the median vein, which is farther from the cubitus vein in ''P. frazieri'' then in ''P. washingtonensis''. In the time after the species description and reexamination of the type specimen by Emerson in 1971 the tip half of fossil has been lost, with only the basal half surviving.


''P. calico''

''P. calico'' was the last North American species of ''Parastylotermes '' to be described, with William D. Pierce publishing the species description in 1958. The isolated wing fragment, ''LACM 533'', from which the species was described was found in 1957 by Wilma Webster in the Calico Mountains,
San Bernardino County, California San Bernardino County (), officially the County of San Bernardino, is a County (United States), county located in the Southern California, southern portion of the U.S. state of California, and is located within the Inland Empire area. As of the ...
. The nodule on which the fossil is preserved is housed in the L.A. County Museum. Due to the incomplete nature of the wing, being only the basal of the wing, it can not be determined if it was a fore or hind-wing though the apparent lack of an anal vein suggests that it may be a fore-wing. While agreeing with Pierce that the description of the wing as a separate species, Emerson noted that there were few features distinguishing ''P. calico'' from ''P. washingtonensis''.


''P. krishnai''

''P. krishnai'' is known from two adults, which are inclusions in amber, making it the second species known from amber inclusions. 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 ...
amber specimen, number ''Tad-277'', is currently housed in the fossil collection of the
Birbal Sahni Birbal Sahni Royal Society, FRS (14 November 1891 – 10 April 1949) was an Indian paleobotanist who studied the fossils of the Indian subcontinent. He also took an interest in geology and archaeology. He founded what is now the Birbal Sahni Ins ...
Institute of Palaeobotany in
Lucknow Lucknow (, ) is the capital and the largest city of the Indian state of Uttar Pradesh and it is also the second largest urban agglomeration in Uttar Pradesh. Lucknow is the administrative headquarters of the eponymous district and division ...
, India and the second known adult, ''Tad-96'' is in the collections of the
American Museum of Natural History The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
. The holotype is composed of a mostly complete adult of indeterminate sex, whereas ''Tad-96'' is a crushed adult, though enough features are present to strongly indicate its placement in ''P. krishnai''. Cambay amber dates to between fifty and fifty-two million years old, placing it in the Early to Mid Ypresian age of the Eocene, and was preserved in a
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
shore environment. The amber formed from a dammar type resin which is produced mainly by trees in the family
Dipterocarpaceae Dipterocarpaceae is a family of 16 genera and about 695 known species of mainly tropical lowland rainforest trees. The family name, from the type genus ''Dipterocarpus'', is derived from Greek (''di'' = two, ''pteron'' = wing and ''karpos'' = fru ...
. The specimens were recovered from the Tadkeshwar lignite mine, located in
Gujarat State Gujarat (, ) is a state along the western coast of India. Its coastline of about is the longest in the country, most of which lies on the Kathiawar peninsula. Gujarat is the fifth-largest Indian state by area, covering some ; and the ninth- ...
, during collecting trips in January 2009 and 2010. The fossils were first studied by paleoentomologists Michael S. Engel and David Grimaldi.and their 2011
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 journal ''
ZooKeys ''ZooKeys'' is a peer-reviewed open access scientific journal covering zoological taxonomy, phylogeny, and biogeography. It was established in 2008 and the editor-in-chief is Terry Erwin (Smithsonian Institution). It is published by Pensoft Publ ...
''. 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 ...
''krishnai'' is in honor of Kumar Krishna, considered a world authority on fossil and living termites. The species has a body length of approximately and a fore-wing length of making it the smallest species known. ''P. krishnai'' also shows a distinct more basal placement of the Medial vein fork then seen in the other species with the wing tip preserved. A notable distinction between ''P. robustus'' and ''P. krishnai'' is difference in numbers of antenna segments. ''P. robustus'' has between sixteen and seventeen segments while ''P. krishnai'' has only fourteen.


References

{{Taxonbar, from=Q7136046 Termite genera Eocene insects Fossil taxa described in 1913 Fossil taxa described in 1931 Fossil taxa described in 1949 Fossil taxa described in 1954 Fossil taxa described in 1959 Fossil taxa described in 2011 Miocene insects Cenozoic insects of Asia Prehistoric insects of North America Prehistoric insects of Europe