Ithonioidea

Ithonidae, commonly called moth lacewings and giant lacewings, is a small family of winged insects of the insect order Neuroptera. The family contains a total of ten living genera, and over a dozen extinct genera described from fossils. The modern Ithonids have a notably disjunct distribution, while the extinct genera had a more global range. The family is considered one of the most primitive living neuropteran families. The family has been expanded twice, first to include the genus '' Rapisma'', formerly placed in the monotypic family Rapismatidae, and then in 2010 to include the genera that had been placed into the family Polystoechotidae. Both Rapismatidae and Polystoechotidae have been shown to nest into Ithonidae ''sensu lato''. The larvae of ithonids are grub-like, subterranean and likely phytophagous (plant feeding).

Description and ecology

Ithonidae are typically medium to large-sized neuropterans. P. S. Welch conducted research in 1914 on "Polystoechotidae" larvae resulting in the conclusion that the larvae were carnivorous. The larvae live a subterranean life. The first instar stage for modern larvae is grub-like, resembling those of scarabaeiform beetles. Robert J. Tillyard first described the larvae of '' Ithone fusca'' in 1922, noting the unusual grub-like shape, similar to fruit-chafers and June beetles, as unique among Neuroptera species. The odd larval shape of Ithionidae larvae has been, and continues to be, a subject of interest among entomologists. Tillyard suggested the larvae to be carnivorous, preying upon Scarabaeidae larvae, attacking with the large mandibles and "sucking them dry". Further study by Faulkner in 1990 has shown the immature Ithonidae to be phytophagous, a condition unique among neuropterans. The adults of at least some ithonids are carnivorous. Mass emergences have been recorded from the genera '' Oliarces'' and '' Ithone'', usually associated with seasonal rains.

Range

Modern Ithonidae genera are distributed in both the Western and Eastern Hemispheres, with three species in the Nearctic realm, and four in the Neotropical realm of the Western Hemisphere. Of the genera in the Eastern Hemisphere, three are found in the Australasian realm and one in Indomalaya. The habitats are varied with the genera falling into two general groups, arid taxa and forest taxa. Those genera found in arid regions are '' Ithone'', '' Varnia'', and'' Oliarces''. Four of the Neotropical species inhabit areas with forested microthermal to lower mesothermal climates. '' Polystoechotes'' has the widest modern range with '' P. punctatus'' inhabiting forests across North America south along the mountains to Panama and '' P. gazullai'' inhabiting forests in Central Chile. The North American range of ''P. punctatus'' has diminished, disappearing from eastern North America by the 1950s, and now appears restricted to Western North America. '' Platystoechotes lineatus'', the sole species in ''Platystoechotes'', is restricted in range to the Sierra Nevada mountains in California, USA. The remaining modern species '' Fontecilla graphicus'' shares habitat with ''P. gazullai'', also inhabiting forests in Central Chile.

Ithonid specimens have been described from fossils dating between the Early Jurassic and the late Eocene, and from sites that indicate a wider geographic range then seen in the modern taxa. Early genera from the Jurassic have been found in China, Kazakhstan, Kyrgyzstan, and Tajikistan. The next grouping of Mesozoic fossils belong to the Cretaceous genus '' Principiala'', which has species described from Brazil, England and tentatively China. The youngest paleorange for the family includes sites in western North America such as the Florissant formation in Colorado and the Okanagan Highland sites in Washington, USA and British Columbia, Canada. Species of the genus '' Palaeopsychops'' have been found in the Eocene Fur Formation of Denmark. The genus '' Elektrithone'', described in 2014, extended the family's fossil range to include Baltic amber.

Systematics and taxonomy

The Ithonidae are regarded as the most primitive living members of Neuroptera. The genus ''Rapisma'' was formerly considered to be the sole genus in the family Rapismatidae, however work in the 1990s and 2000s resulted in the inclusion of ''Rapisma'' in Ithonidae. The history of Ithonidae and Polysteochotidae is more complex, with several genera having been transferred between each other over the years. Phylogenetic analysis and associated genetic sequencing of three genetic markers for the two families was published in 2010. The results of the analysis showed a clear grouping of the three Australian genera ''Ithone'', ''Megalithone'' and ''Varnia'' in the family Ithonidae, while all the remaining genera are nested into an expanded Polysteochotidae, given the overall interrelation between the groups, the authors opted to combine the two into an expanded single family Ithonidae.

Ithonidae forms a sister group to the remaining families in the suborder Hemerobiiformia. Ithonidae includes a number of extinct genera described from fossils dating between the Early Jurassic and the Late Eocene. The genera of Ithonidae are clustered into three informal groups:

The Ithonid group:
*†'' Guithone'' Middle Jurassic (Callovian), Daohugou, China
*'' Ithone'' Australia
*'' Megalithone'' Australia
*'' Varnia'' Australia

The Polystoechotid group:
*†'' Elektrithone'' (Eocene, Baltic Amber)
*'' Fontecilla'' Chile
*†'' Frustumopsychops'' Late Jurassic (Tithonian), Shar-Teg, Mongolia
*†'' Jurapolystoechotes'' Middle Jurassic (Callovian) Daohugou Bed, China
*†'' Kirgisella'' Karabastau Formation, Late Jurassic ( Oxfordian/Kimmeridgian), Kazakhstan
*†'' Lasiosmylus'' Early Cretaceous (Aptian), Yixian Formation, China
*†'' Mesopolystoechus'' Early Jurassic (Toarcian), Sulyukta Formation, Kyrgyzstan; Callovian, Jiulongshan Formation, China
*†'' Osmyloides'' (Middle/Late Jurassic Callovian/ Oxfordian, Karabastau Formation, Kazakhstan)
*†'' Palaeopsychops'' Eocene (Ypresian), Coldwater Beds, Canada, Fur Formation, Denmark, Klondike Mountain Formation, USA
*†'' Paleopterocalla'' (Callovian/Oxfordian, Karabastau Formation, Kazakhstan)
*†'' Panfilovdvia'' (Callovian/Oxfordian, Karabastau Formation, Kazakhstan)
*'' Platystoechotes'' Western North America
*†'' Polystoechotites'' (Eocene, Coldwater Beds, Canada, Klondike Mountain Formation, Florissant Formation, USA
*'' Polystoechotes'' North America, Chile
*†'' Puripolystoechotes'' (Daohugou, China, Callovian)
*†'' Ricaniella'' Eocene (Ypresian), Allenby Formation, Canada

The Rapismatid Group:
*†'' Allorapisma'' Eocene (Ypresian), Klondike Mountain Formation, Washington
*'' Adamsiana'' Central America
*†'' Burmithone'' Late Cretaceous (Cenomanian), Burmese amber, Myanmar
*'' Narodona'' Mexico
*'' Oliarces'' Western North America
*†'' Principiala'' Early Cretaceous (Barremian), Weald Clay, United Kingdom, Aptian Crato Formation, Brazil
*†'' Sinuijuala'' Early Cretaceous (Aptian) Sinuiju Formation, North Korea
*'' Rapisma'' South, Southeast and Eastern Asia

References

{{Taxonbar, from=Q1938392

Neuroptera
Neuroptera families
Taxa named by Edward Newman