Alienopteridae
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Alienopteridae is an extinct family of dictyopterans, known from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
to the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
epoch. They are noted for their unusual combination of features not found in other dictyopterans.


Description

Alienopterid nymphs vary in morphology, some are superficially ant-like, while others are more robust. In adult alienopterids, the first set of wings are reduced into shortened sclerotised scale-like structures. The compound eyes are large, and the head generally triangular shaped and able to rotate.


Taxonomy

It was originally assigned to its own order Alienoptera by Bai et al., 2016. It was reassigned to the dictyopteran superfamily Umenocoleoidea as sister family to the beetle-like Umenocoleidae by Vršanský ''et al.'' (2018), and a more recent analysis similarly places Alienopteridae and Umenocoleidae as sister taxa within Dictyoptera, but placing both lineages outside of
Blattodea Blattodea is an order (biology), order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetics, genetic and molecular evidence suggests they evolved from within the cockroach ...
. 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 (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 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
(latest
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
-earliest
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
~ 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. Th ...
found in
Myanmar Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and has ...
; though an additional two genera ('' Apiblatta'' and '' Vcelesvab'') are from the late
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
Crato Formation (
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
), and the two youngest genera ('' Chimaeroblattina'' and '' Grant'') are from the middle
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
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 (Colorado River), Green River in Colorado, Wyoming, and Utah. The sedimen ...
(
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
).


Ecology

In a 2018 paper, some alienopterid adults were suggested as mimics of various hymenopterans, including bees and
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s. However this was mostly rejected in another study as lacking evidence. A 2021 study alternatively suggested that some alienopterid nymphs functioned 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 pollen (probably from a cycad 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 carpel, 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 ...
. Alienopterid adults have been suggested to be predators adapted to moving through dense foliage, with the small sclerotised forewings being an adaptation for flight capability.


Genera

*Superfamily † Umenocoleoidea Chen & Tan, 1973 **Family †Alienopteridae Bai ''et al''., 2016 *** †'' Aethiocarenus'' Poinar & Brown, 2016 *** †'' Alienopterella'' Kočárek, 2018 *** †'' Alienopterus'' Bai ''et al''. 2016 *** †'' Apiblatta'' Barna & Bigalk, 2018 *** †'' Caputoraptor'' Bai ''et al''., 2018 *** †'' Chimaeroblattina'' Barna, 2018 *** †'' Formicamendax'' Hinkelman, 2019 *** †'' Grant'' 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 International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
aged Orapa kimberlite pipe sediments in
Botswana Botswana, officially the Republic of Botswana, is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory part of the Kalahari Desert. It is bordered by South Africa to the sou ...
. 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