Ithonidae

File:Allorapisma chuorum Holotype SR 08-14-01.jpg holotype]]

Ithonidae are typically medium to large-sized neuropterans.{{cite journal |last = Archibald |first = S.B. |author2 = Makarkin V.N. |title = Tertiary Giant Lacewings (Neuroptera: Polystechotidae): Revision and Description of New Taxa From Western North America and Denmark |journal = Journal of Systematic Palaeontology |volume = 4 |issue = 2 |pages = 119–155 |year = 2006 |url = http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=438987 |access-date = January 27, 2010 |doi = 10.1017/S1477201906001817 |s2cid = 55970660 |archive-date = June 4, 2011 |archive-url = https://web.archive.org/web/20110604115043/http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=438987 |url-status = dead }} P. S. Welch conducted research in 1914 on "Polystoechotidae" larvae resulting in the conclusion that the larvae were carnivorous.{{cite journal |last = Grebennikov |first = V. |title = Grub-like larvae of Neuroptera (Insecta): a morphological review of the families Ithonidae and Polystoechotidae and a description of Oliarces clara |journal = European Journal of Entomology |volume= 101 |pages= 409–417 |year= 2004 |issue = 3 |url= https://www.eje.cz/pdfarticles/721/eje_101_3_409_IthonGreb.pdf |access-date=January 27, 2010 |doi=10.14411/eje.2004.056|doi-access= free }} The larvae live a subterranean life.{{Cite journal |last1=Li |first1=Di |last2=Friedrich |first2=Frank |last3=Jandausch |first3=Kenny |last4=Pohl |first4=Hans |last5=Liu |first5=Xingyue |last6=Beutel |first6=Rolf G. |date=October 2022 |title=Unearthing underground predators: The head morphology of larvae of the moth lacewing genus Ithone Newman (Neuroptera: Ithonidae) and its functional and phylogenetic implications |url=https://onlinelibrary.wiley.com/doi/10.1111/syen.12556 |journal=Systematic Entomology |language=en |volume=47 |issue=4 |pages=618–636 |doi=10.1111/syen.12556 |s2cid=249359273 |issn=0307-6970}} 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.{{Cite journal |last=De Jong |first=Grant D. |date=July 2011 |title=Observations on the Biology of Polystoechotes punctatus (Fabricius) (Neuroptera: Ithonidae): Adult Trophic Status, Description of the Male Reproductive System, and Associations with Mites |url=http://www.bioone.org/doi/abs/10.4289/0013-8797.113.3.291 |journal=Proceedings of the Entomological Society of Washington |language=en |volume=113 |issue=3 |pages=291–298 |doi=10.4289/0013-8797.113.3.291 |s2cid=198148984 |issn=0013-8797}} Mass emergences have been recorded from the genera Oliarces and Ithone, usually associated with seasonal rains.{{cite journal |last1=Winterton |first1=SL |last2=Makarkin |first2=VN |year=2010 |title=Phylogeny of Moth Lacewings and Giant Lacewings (Neuroptera: Ithonidae, Polystoechotidae) Using DNA Sequence Data, Morphology, and Fossils |journal=Annals of the Entomological Society of America |volume=103 |issue=4 |pages=511–522 |doi=10.1603/an10026|s2cid=49384430 |doi-access=free }}

File:Palaeopsychops marringerae Holotype SR 97-08-05 v2.jpg fossil, 50 million years old, Washington, USA]]

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,{{Cite book |author=S. Marshall |title=Insects: Their Natural History And Diversity: With a Photographic Guide to Insects of Eastern North America |publisher=Firefly Books |year=2006 |isbn=978-1-55297-900-6}} 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.{{Cite journal|last1=Makarkin |first1=VN |last2=Wedmann |first2=S |last3=Weiterschan |first3=T |year=2014 |title=First record of the family Ithonidae (Neuroptera) from Baltic amber |journal=Zootaxa |volume=3796 |issue=2 |pages=385–393 |doi=10.11646/zootaxa.3796.2.10 |pmid=24870683 }}

File:Puripolystoechotes pumilus holotype fig1a.png,
Jiulongshan Formation ]]

File:Elektrithone expectata holotype Baltic amber Fig1.png
Baltic amber]]

File:Ricaniella antiquata Scudder 1895 pl1 Fig3.png
Allenby Formation
(1895 illustration)]]

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 {{small|Zheng, Ren, & wang, 2016}} Middle Jurassic (Callovian), Daohugou, China
• Ithone {{small|Newman, 1838}} Australia
• Megalithone {{small|Riek, 1974}} Australia
• Varnia {{small|Walker, 1860}} Australia

The Polystoechotid group:

• †Elektrithone {{small|Makarkin, Wedmann, & Weiterschan, 2014}} (Eocene, Baltic Amber)
• Fontecilla {{small|Navás, 1931}} Chile
• †Frustumopsychops {{small|Khramov 2014}} Late Jurassic (Tithonian), Shar-Teg, Mongolia
• †Jurapolystoechotes {{small|Ren, Engel, & Lü, 2002}} Middle Jurassic (Callovian) Daohugou Bed, China
• †Kirgisella {{small|Martynov, 1925}} Karabastau Formation, Late Jurassic (Oxfordian/Kimmeridgian), Kazakhstan
• †Lasiosmylus {{small|Ren & Guo, 1996}} Early Cretaceous (Aptian), Yixian Formation, China
• †Mesopolystoechus {{small|Martynov, 1937}} Early Jurassic (Toarcian), Sulyukta Formation, Kyrgyzstan; Callovian, Jiulongshan Formation, China
• †Osmyloides {{small|Panfilov, 1980}} (Middle/Late Jurassic Callovian/Oxfordian, Karabastau Formation, Kazakhstan)
• †Palaeopsychops {{small|Anderson, 2001}} Eocene (Ypresian), Coldwater Beds, Canada, Fur Formation, Denmark, Klondike Mountain Formation, USA
• †Paleopterocalla {{small|(Panfilov, 1980)}} (Callovian/Oxfordian, Karabastau Formation, Kazakhstan)
• †Panfilovdvia {{small|(Panfilov, 1980)}} (Callovian/Oxfordian, Karabastau Formation, Kazakhstan)
• Platystoechotes {{small|Carpenter, 1940}} Western North America
• †Polystoechotites {{small|Archibald & Makarkin, 2006}} (Eocene, Coldwater Beds, Canada, Klondike Mountain Formation, Florissant Formation, USA
• Polystoechotes {{small|Burmeister, 1839}} North America, Chile
• †Puripolystoechotes {{small|Yang et al. 2019}} (Daohugou, China, Callovian)
• †Ricaniella {{small|Meunier, 1897}} Eocene (Ypresian), Allenby Formation, Canada

The Rapismatid Group:

• †Allorapisma {{small|Makarkin & Archibald, 2009}} Eocene (Ypresian), Klondike Mountain Formation, Washington
• Adamsiana {{small|Penny, 1996}} Central America
• †Burmithone {{small|Lu et al., 2017}} Late Cretaceous (Cenomanian), Burmese amber, Myanmar
• Narodona {{small|Navás, 1929}} Mexico
• Oliarces {{small|Banks, 1908}} Western North America
• †Principiala {{small|Makarkin & Menon, 2007}} Early Cretaceous (Barremian), Weald Clay, United Kingdom, Aptian Crato Formation, Brazil
• †Sinuijuala {{small|So & Won, 2022}} Early Cretaceous (Aptian) Sinuiju Formation, North Korea
• Rapisma {{small|McLachlan, 1866}} South, Southeast and Eastern Asia

{{clade

|1={{clade

|label1=Polysteochotidae

|1={{clade

|1={{clade

|1=Adamsiana

|2={{clade

|1={{clade

|1=†Allorapisma

|2=†Principiala

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|2=Rapisma

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|3=Oliarces

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|2=Platystoechotes

|3={{clade

|1=Fontecilla

|2=Polystoechotes

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}}

|label2=Ithonidae

|2={{clade

|1=Ithone

|2=Megalithone

|3=Varnia

}}

}}

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{{Neuroptera|2}}

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Category:Neuroptera

Category:Neuroptera families

Category:Taxa named by Edward Newman