Alienopteridae
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Alienopteridae is an extinct family of
dictyoptera Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the o ...
ns, known from the Mid-Cretaceous to
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
. They are noted for their unusual combination of features not found in other dictyopterans.


Taxonomy

It was originally assigned to its own order Alienoptera by Bai et al., 2016. It was reassigned to the dictyopteran superfamily
Umenocoleoidea Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the or ...
as sister family to the beetle-like
Umenocoleidae Umenocoleidae is an extinct family of dictyopteran insects known from the Cretaceous. They are considered to be closely related to the Alienopteridae. They were originally considered to be beetles due to their beetle-like morphology, with sclero ...
by Vršanský ''et al.'' (2018), and a more recent analysis similarly places Alienopteridae and
Umenocoleidae Umenocoleidae is an extinct family of dictyopteran insects known from the Cretaceous. They are considered to be closely related to the Alienopteridae. They were originally considered to be beetles due to their beetle-like morphology, with sclero ...
as sister taxa within
Dictyoptera Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the o ...
, but placing both lineages outside of
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 suggests they evolved from within the cockroach lineage, cladistically ...
. A 2021 study revived the order Alienoptera for the clade containing Alienopteridae and Umenocoleidae, with a cladistic analysis placing Alienoptera as the sister clade to
Mantodea Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
(praying mantises).


Distribution

The majority of the alienopterid genera are known from the mid
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
(latest
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 M ...
-earliest
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
~ 100 million years ago)
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
found in
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
; though an additional two genera ('' Apiblatta'' and '' Vcelesvab'') 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 ...
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous ( Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly duri ...
(
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
), and the two youngest genera ('' Chimaeroblattina'' and ''
Grant Grant or Grants may refer to: Places *Grant County (disambiguation) Australia * Grant, Queensland, a locality in the Barcaldine Region, Queensland, Australia United Kingdom *Castle Grant United States * Grant, Alabama *Grant, Inyo County, C ...
'') are from the middle
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very fine ...
(
Colorado Colorado (, other variants) is a state in the 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 western edge of t ...
,
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
).


Ecology

In a 2018 paper, some alienopterid adults were suggested as
mimics Materialise Mimics is an image processing software for 3D design and modeling, developed by Materialise NV, a Belgian company specialized in additive manufacturing software and technology for medical, dental and additive manufacturing industries ...
of various
hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
ns, including bees and
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s. However this was mostly rejected in another study as lacking evidence. A 2021 study alternatively suggested that some alienopterid nymphs functoned as ant mimics, based on morphological features that closely resembled contemporary sphecomyrmine ants, and suggested that the adult genus '' Teyia'' was also a wasp mimic, though mimicry in other adult alienopterids was rejected. One alienopterid nymph was found laden with
gymnosperm 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 ...
pollen (probably from a
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
or a bennettitalean), suggesting that it consumed pollen as part of its diet, and that it also acted as a
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
. Alienopterid adults were probably predators adapted to moving through dense foliage, with the small
sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qui ...
forewings being an adaptation for flight capaciity.


Genera

*Superfamily †
Umenocoleoidea Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the or ...
Chen & Tan, 1973 **Family †Alienopteridae Bai ''et al''., 2016 *** †'' Aethiocarenus'' Poinar & Brown, 2016 *** †'' Alienopterella'' Kočárek, 2018 *** †''
Alienopterus ''Alienopterus brachyelytrus'' is an extinct insect described from a 99 million year old fossil found in Burmese amber from the Hukawng Valley of Myanmar. It was the first known member of the order Alienoptera until 2018, when the second and th ...
'' Bai ''et al''. 2016 *** †'' Apiblatta'' Barna & Bigalk, 2018 *** †''
Caputoraptor ''Caputoraptor'' is an extinct genus of insect from the Cenomanian aged Burmese amber containing two species: the type species ''Caputoraptor elegans'', as well as ''Caputoraptor vidit''. It is part of the extinct order Alienoptera. ''C. elegans ...
'' Bai ''et al''., 2018 *** †'' Chimaeroblattina'' Barna, 2018 *** †'' Formicamendax'' Hinkelman, 2019 *** †''
Grant Grant or Grants may refer to: Places *Grant County (disambiguation) Australia * Grant, Queensland, a locality in the Barcaldine Region, Queensland, Australia United Kingdom *Castle Grant United States * Grant, Alabama *Grant, Inyo County, C ...
'' Aristov, 2018 *** †'' Meilia'' Vršanský & Wang, 2018 *** †'' Teyia'' Vršanský ''et al''., 2018 *** †'' Vcelesvab'' Vršanský et al. 2018 An undescribed species is also known from the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
aged
Orapa Orapa is a town located in the Central District of Botswana. It is the site of the Orapa diamond mine, the largest diamond-producing mine in the world, and is considered to be the diamond capital of the country. Nearby is another kimberlite min ...
kimberlite pipe Volcanic pipes or volcanic conduits are subterranean geology, geological structures formed by the violent, supersonic eruption of deep-origin volcanoes. They are considered to be a type of ''diatreme''. Volcanic pipes are composed of a deep, narro ...
sediments in
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label=Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kalahar ...
. Luo, Xu & Jarzembowski (2020) transferred ''Alienopterix'' and ''Vzrkadlenie''s, originally described as alienopterids, to the family Cratovitismidae. A 2021 study later considered ''Alienopterix'' an umenocoleid instead.


References

Prehistoric insect families Dictyoptera {{Paleo-insect-stub