Palaeontinidae, commonly known as giant cicadas, 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 ...
family of
cicadomorphs. They existed during the
Mesozoic era
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising ...
of
Europe,
Asia, and
South America.
The family contains around 30 to 40
genera
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 nomenclat ...
and around a hundred
species.
Discovery
The first palaeontinid discovered was ''
Palaeontina oolitica''. It consisted of a single
forewing collected from the
Taynton Limestone Formation (Stonesfield Slate) of
Oxfordshire
Oxfordshire is a ceremonial and non-metropolitan county in the north west of South East England. It is a mainly rural county, with its largest settlement being the city of Oxford. The county is a centre of research and development, primarily ...
,
England by the
English natural historian Edward Charlesworth. It was first described in 1873 by the English
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 ...
Arthur Gardiner Butler in his book ''Lepidoptera Exotica; or, Descriptions and Illustrations of Exotic Lepidoptera''. Butler claimed that it was the oldest
butterfly ever recovered, having mistakenly identified it as a butterfly of the family
Nymphalidae.
Description and paleobiology
Palaeontinids had large bodies covered with bristles (
setae). They had small heads and broad wings. They superficially resemble moths.
Large palaeontinids like ''
Colossocossus'' had forewings that reached the length of .
They possessed an inflated
frons and a long
rostrum (piercing and sucking mouthpart), indicating that they fed on
xylem fluids like some other modern
hemiptera
Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
ns.
Some authors have proposed that the
host plants of palaeontinids to be
ginkgophyte
Ginkgoales are a gymnosperm order containing only one extant species: ''Ginkgo biloba'', the ginkgo tree. It is monotypic, (the only taxon) within the class Ginkgoopsida, which itself is monotypic within the division Ginkgophyta . The order inc ...
s based on the geographic distribution of both groups, however other authors have argued that this association is likely to be spurious, given that that paleontinids also occur in areas with no ginkgophytes. Some authors have suggested that the decline of
gymnosperms
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
and the rise of
angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s (flowering plants) during the Cretaceous could have been a factor in their extinction.
Numerous newly evolved
insectivorous
A robber fly eating a hoverfly
An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects.
The first vertebrate insectivores were ...
animals (feathered
theropods, primitive
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s, and early
birds) may have also contributed significantly to their extinction.
Most species of palaeontinids exhibit
cryptic coloration
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
.
The patterns on their wings protected them as they perched on branches and fed on sap. They may also have served as
secondary sexual characteristics
Secondary sex characteristics are features that appear during puberty in humans, and at sexual maturity in other animals. These characteristics are particularly evident in the sexually dimorphic phenotypic traits that distinguish the sexes of a sp ...
. The color patterns can vary slightly within the same species.
Palaeontinids, like modern cicadas, possess four membranous wings supported by veins. The length and width ratio of the wings can vary within the same species, sometimes as a result of fossil preservation.
Early Jurassic palaeontinids, like ''
Suljuktocossus'', exhibit the most primitive wing forms in the family.
The forewing was elliptical with the "nodal line" (the area where the wing bends during flight, also known as the "transverse flexion line") more or less dissecting through the center of the wing. The hindwing was short and broad. The bases of the forewings overlapped that of the hindwings like in modern butterflies. Taken together with their large bodies, these characteristics indicate that they were fast but moderately versatile fliers.
In contrast, later palaeontinids like the Upper Jurassic ''
Eocicada'' and Early Cretaceous ''
Ilerdocossus'' had triangular forewings with the flexion line closer to the base. They had smaller and narrower hindwings that did not overlap with the forewing. These indicate that they were highly versatile fliers, able to fly with a wide range of speeds and agility like modern
wasps and
sphinx moths.
They also possessed changes to the leading edge of their forewings, suggesting an overall gain in
lift.
The trend of forewing elongation is most evident in members of the family
Mesogereonidae, an early offshoot and close relatives of palaeontinids.
Classification
The family was first erected by the
Austrian
Austrian may refer to:
* Austrians, someone from Austria or of Austrian descent
** Someone who is considered an Austrian citizen, see Austrian nationality law
* Austrian German dialect
* Something associated with the country Austria, for example: ...
entomologist
Anton Handlirsch
Anton Handlirsch or Anton Peter Josef Handlirsch (20 January 1865, Vienna – 28 August 1935, Vienna) was an Austrian entomologist. He worked on many groups including Hemiptera, Hymenoptera and Neuroptera. His most significant work was in the stu ...
in 1908. Like Butler, Handlirsch insisted that palaeontinids were members of
lepidoptera
Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
n
Heteroneura (butterflies and moths). Palaeontinids were then only known mostly from poorly preserved specimens like ''
Palaeontina'' and ''
Eocicada''. He claimed they were related to the extant family
Limacodidae (slug moths).
The English entomologist
Edward Meyrick supported the lepidopteran conclusion, though he believed they belonged to the family
Hepialidae (ghost moths) instead. He said "There is little doubt that it
.e. ''Palaeontina oolitica''belongs to the Hepialidae."
The
Belgian entomologist
Auguste Lameere challenged this conclusion, claiming palaeontinids were more closely related to the extant family
Cicadidae (cicadas). The English-
Australian entomologist and
geologist Robert John Tillyard supported Lameere's conclusion, noting that the wings of palaeontinid fossils lacked the characteristic scales of lepidopterans but instead had tubercules, pits, and cross-ridges like those found in modern cicadas.
He also cited characteristics of wing venation that distinctly differs from that of lepidopterans.
Palaeontinidae are currently classified under the extinct superfamily
Palaeontinoidea
Palaeontinoidea is an extinct superfamily of cicadomorph hemipteran insects. This superfamily contains three families.
Description
Palaeontinoids were comparatively large, cicada-like insects that existed from the Upper Permian to the Middle ...
along with the families
Dunstaniidae and
Mesogereonidae.
They are classified under
infraorder Cicadomorpha of the
hemiptera
Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
ns (true bugs).
The name Cicadomorphidae was once proposed as a replacement for the name Palaeontinidae in 1956 by the Australian entomologist J.W. Evans. This was because of Handlirsch's earlier insistence that the
type species ''Palaeontina oolitica'' may not have been
Hemiptera
Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
n. However, Evans later conceded that retaining the name Palaeontinidae was preferable as the drawings Handlirsch based his conclusions on were from badly preserved specimens.
Evolution
Riek (1976) originally considered Palaeontinoidea to be the descendants of the family Cicadoprosbolidae (currently known as the family
Tettigarctidae), insects believed to be transitional between the ancestral cicada-like family
Prosbolidae and the modern family
Cicadidae.
Wang ''et al'' (2009), however, notes that palaeontinoids more closely resemble prosbolids in agreement with earlier studies by Wootton (1971), Shcherbakov (1984), and Shcherbakov and Popov (2002). They conclude that palaeontinoids descended directly from the family Prosbolidae rather than from tettigarctids.
Modern cicadas therefore, did not descend directly from Palaeontinidae.
Within Palaeontinoidea, the family Dunstaniidae (
Upper Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
to
Lower Jurassic of
Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
South Africa, and
China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
) is ancestral to palaeontinids. Both are distinct from the only other member of the superfamily, the more primitive and specialized family Mesogereonidae (Upper Triassic of Australia and South Africa).
Distribution and geologic time range
The oldest known member of the group is ''
Hallakkungis ''from South Korea dating to the
Norian
The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.
Stratigraphic defi ...
stage of the
Late Triassic (''ca''. 227 – ''ca''. 208.5 Mya)
and the youngest members are from the late
Aptian
The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
age of the
Lower Cretaceous (~115-113 Mya).
They achieved their greatest diversity during the
Jurassic period.
Palaeontinid fossils are abundant in
Eurasia and
South America.
Fossils have been recorded in
Brazil,
China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,
Russia,
Germany, the
Transbaikal region,
Tajikistan,
Turkmenistan,
Kyrgyzstan,
Kazakhstan,
Spain, and the
United Kingdom. Important
localities for palaeontinid fossils include the
Crato Formation Lagerstätte of
Brazil and the
Yixian Formation,
Haifanggou (or Jiulongshan) Formation, and the
Daohugou Beds of
China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
.
Genera
The following is the list of
genera
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 nomenclat ...
classified under Palaeontinidae:
*''
Abrocossus''
Wang & Zhang, 2007 ''in'' Wang et al. 2007a -
Daohugou Beds, Middle Jurassic, East Asia
*''
Archipsyche''
Handlirsch, 1906–1908 -
Solnhofen Formation Late Jurassic, Central Europe
*''
Asiocossus ''
Becker-Migdisova, 1962 -
Dzhil Formation Dzhil may refer to:
* Jil, Armenia
* Cil, Azerbaijan
Cil (also, Kazımlı Cil and Dzhil’) is a village and municipality in the Lankaran Rayon of Azerbaijan. It had a population of 2,279 as of 2007.
Notable natives
* Allahshukur Pashazadeh, ...
, Early Jurassic, Central Asia
*''
Baeocossus ''
Menon et al. 2005 -
Crato Formation Early Cretaceous, Eastern South America
*''
Cicadomorpha ''
Martynov, 1926 -
Karabastau Formation, Late Jurassic, Central Asia;
Glushkovo Formation Glushkovo (russian: Глушково) is the name of several inhabited localities in Russia.
;Urban localities
*Glushkovo, Glushkovsky District, Kursk Oblast, a work settlement in Glushkovsky District of Kursk Oblast
;Rural localities
* Glushko ...
,
Ukurei Formation, Late Jurassic North Asia
*''
Colossocossus ''
Menon et al. 2005 - Crato Formation, Early Cretaceous, Eastern South America
*''
Cratocossus ''
Martins-Neto, 1998 - Crato Formation, Early Cretaceous, Eastern South America
*''
Daohugoucossus ''
Wang et al. 2006b - Daohugou, Middle Jurassic, East Asia
*''
Eocicada ''
Oppenheim, 1888 - Solnhofen Formation, Late Jurassic, Central Europe
*''
Eoiocossus ''
Wang et al. 2006c ''in'' Wang et al. 2006c - (includes ''
Papilioncossus''
Wang et al. 2007c) Daohugou Middle Jurassic, East Asia
*''
Gansucossus ''
Wang et al. 2006b -
Dashankou Formation,
Jiulongshan Formation
The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China.
The formation consists of coarse conglomerates ...
, Daohugou Middle Jurassic, East Asia
*''
Hallakkungis ''
Nam, Wang, & Szwedo, 2017 -
Amisan Formation Upper Triassic, South Korea
*''
Hamicossus ''
Wang & Ren 2007ab - Daohugou Middle Jurassic, East Asia
*''
Ilerdocossus ''
Gomez-Pallerola, 1984 - (includes ''
Wonnacottella ''
Whalley & Jarzembowski, 1985 and ''
Liaocossus ''
Ren et al., 1998)
La Pedrera de Rúbies Formation
The La Pedrera de Rúbies Formation is an Early Cretaceous ( late Berriasian to early Barremian geologic formation in Catalonia, Spain. The formation crops out in the area of the Montsec in the Organyà Basin. At the La Pedrera de Meià localit ...
,
Weald Clay
Weald Clay or the Weald Clay Formation is a Lower Cretaceous sedimentary rock unit underlying areas of South East England, between the North and South Downs, in an area called the Weald Basin. It is the uppermost unit of the Wealden Group of ro ...
,
Yixian Formation Early Cretaceous, Western Europe & East Asia
*''
Limacodites ''
Handlirsch, 1906–1908 - Solnhofen Formation, Late Jurassic, Central Europe
*''
Martynovocossus ''
Wang & Zhang 2008 ''in'' Wang et al. 2008 - (= ''
Pseudocossus
''Pseudocossus'' is a genus of moths of the family Cossidae from Madagascar.afromoths.net
''Type species:'' ''Ps ...
''
Martynov, 1931)
Badaowan Formation, Daohugou,
Kushmurun Formation
Kushmurun ( kk, Құсмұрын; russian: Кушмурун) is a brackish lake in the Kostanay Region, Kazakhstan.
Kushmurun is one of largest natural waterbodies of Kazakhstan. It lies in the northern sector of the Turgay Basin, at the border be ...
,
Cheremkhovskaya Formation
Cheremkhovsky District (russian: Черемхо́вский райо́н) is an administrative district, one of the thirty-three in Irkutsk Oblast, Russia.Charter of Irkutsk Oblast, Article 13 Municipally, it is incorporated as Cheremkhovsk ...
, Early to Middle Jurassic, North Asia; Late Jurassic, Central Asia
*''
Miracossus ''
Ren et al. 1998 - Yixian Formation, Early Cretaceous, East Asia
*''
Montsecocossus ''
Gomez-Pallerola, 1984 - La Pedrera de Rúbies Formation Early Cretaceous, Western Europe
*''
Neimenggucossus ''
Wang & Zhang, 2007 ''in'' Wang et al. 2007a - Daohugou, Middle Jurassic, East Asia
*''
Ningchengia ''
Wang, Zhang & Szwedo, 2009 - (= ''
Fletcheriana''
Evans, 1956 ''in partim'') Daohugou Middle Jurassic, East Asia
*''
Pachypsyche ''
Handlirsch 1906 - La Pedrera de Rúbies Formation Early Cretaceous, Western Europe
*''
Palaeocossus ''
Oppenheim, 1885 - Sagul Formation, Cheremkhovskaya Formation Early Jurassic, Central and North Asia
*''
Palaeontina ''
Butler, 1873 -
Taynton Limestone Formation Middle Jurassic, Western Europe
*''
Palaeontinodes ''
Martynov, 1937 - (= ''
Ijacossus ''
Becker-Migdisova, 1950; Shcherbakov 1985)
Sulyukta Formation,
Sagul Formation, Haifanggou Formation, Cheremkhovskaya Formation, Early to Middle Jurassic, Central and North Asia; Middle Jurassic, Central Asia
*''
Parawonnacottella ''
Ueda, 1997 - Crato Formation, Early Cretaceous, Eastern South America
*''
Phragmatoecicossus ''
Becker-Migdisova, 1949 - Sagul Formation, Early to Middle Jurassic, Central Asia
*''
Phragmatoecites ''
Oppenheim, 1885 - Cheremkhovskaya Formation, Early to Middle Jurassic, North Asia
*''
Plachutella ''
Becker-Migdisova, 1949 - Badaowan Formation, Jiulongshan Formation, Daohugou, Karabastau Formation, Sagul Formation, Cheremkhovskaya Formation, Early to Late Jurassic, Central and Eastern Asia
*''
Prolystra ''
Oppenheim, 1888 syn ''
Beloptesis ''- Solnhofen Formation, Late Jurassic, Central Europe
*''
Protopsyche ''
Handlirsch, 1906–1908 - Solnhofen Formation, Late Jurassic, Central Europe
*''
Shurabocossus ''
Becker-Migdisova, 1949 - Sagul Formation, Early to Middle Jurassic, Central Asia
*''
Sinopalaeocossus ''
Hong, 1983- (= ''
Quadraticossus''
Wang & Ren, 2007a) Daohugou, Haifanggou Formation, Middle Jurassic, East Asia
*''
Suljuktaja ''
Becker-Migdisova, 1949 - Sulyukta Formation, Early to Middle Jurassic, Central Asia
*''
Suljuktocossus ''
Becker-Migdisova, 1949 - (= ''
Fletcheriana''
Evans, 1956 ''in partim'') Sulyukta Formation, Early Jurassic, Central Asia; Daohugou Middle Jurassic, North Asia
*†''
Synapocossus''
Wang et al. 2013 Daohugou Middle Jurassic, East Asia
*†''
Talbragarocossus''
Chen et al. 2019 Talbragar Fossil Fish Bed, Australia, Late Jurassic
*''
Turgaiella ''
Becker-Migdisova & Wootton, 1965 -
Kushmurun Formation
Kushmurun ( kk, Құсмұрын; russian: Кушмурун) is a brackish lake in the Kostanay Region, Kazakhstan.
Kushmurun is one of largest natural waterbodies of Kazakhstan. It lies in the northern sector of the Turgay Basin, at the border be ...
, Jurassic, Central Asia
*''
Valdicossus''
Wang, Zhang & Jarzembowski 2008 - Weald Clay, Early Cretaceous, Western Europe
*''
Yanocossus ''
Ren, 1995 - Yixian Formation, Early Cretaceous, East Asia
*''
Cyllonium ''
Westwood, 1854 - Early Cretaceous, Western Europe (too poorly preserved)
*''
Palaeontinopsis ''
Martynov, 1937 - Early Jurassic, Central Asia (''
nomen dubium
In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application.
Zoology
In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'')
*''
Palaeontinopsis sinensis ''
- Hong, 1986; Zhang, 1997 Middle Jurassic, East Asia
*''
Fletcheriana jurassica ''
- Zhang, 1997
*''
Fletcheriana magna ''
- Riek, 1976
See also
*
Prehistoric Lepidoptera
Prehistoric Lepidoptera are both butterflies and moths that lived before recorded history. The fossil record for Lepidoptera is lacking in comparison to other winged species, and tending not to be as common as some other insects in the habitats t ...
*
Prehistoric insects
Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use of ...
References
External links
*
*
{{Taxonbar, from=Q7126461
Extinct Hemiptera
Triassic insects
Jurassic insects
Cretaceous insects
Prehistoric insect families
Rhaetian first appearances
Aptian extinctions
Taxa named by Anton Handlirsch