Berothidae

Berothidae Temporal range: Late Jurassic–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Spermophorella sp.
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Neuroptera
Clade:	Euneuroptera
Superfamily:	Mantispoidea
Family:	Berothidae Handlirsch, 1906
Subfamilies
See text

The Berothidae are a family of winged insects of the order Neuroptera. They are known commonly as the beaded lacewings.[1] The family was first named by Anton Handlirsch in 1906.[2] The family consists of 24 genera and 110 living species distributed discontinuously worldwide, mostly in tropical and subtropical regions.[3] Numerous extinct species have also been described. Their ecology is poorly known, but in the species where larval stages have been documented, the larvae are predators of termites.[4]

Systematics

Typical posture in life

A considerable fossil diversity of beaded lacewings is known from the Late Jurassic onwards, containing numerous genera which are likewise basal or incertae sedis.

Subfamily Berothimerobiinae

• Berothimerobius Monserrat & Deretsky, 1999

Subfamily Berothinae

• Asadeteva Aspöck & Aspöck, 1981
• Barrowiella Smithers, 1984
• Berlekrumyia Aspöck & Aspöck, 1988
• Berotha Walker, 1860
• †Elektroberotha Makarkin & Ohl 2015 Eocene (Lutetian); Baltic amber, northern Europe
• Isoscelipteron Costa, 1863
• Lekrugeria Navás, 1929
• Lomamyia Banks, 1904
• Nodalla Navás, 1926
• Podallea Navás, 1936
• Quasispermophorella Aspöck & Aspöck, 1986
• Spermophorella Tillyard, 1916
• Spiroberotha Adams, 1990
• Stenobiella Tillyard, 1916
• †Xenoberotha Makarkin 2017 Green River Formation, United States, Eocene

Subfamily Cyrenoberothinae[5]

• †Aggregataberotha Wang et al., 2023 (Cenomanian, Burmese amber, Myanmar)
• †Araripeberotha Martins-Neto & Vulcano 1990 Cretaceous (Aptian); Crato Formation, Brazil
• †Caririberotha Martins-Neto & Vulcano 1990 Cretaceous (Aptian); Crato Formation, Brazil
• Cyrenoberotha MacLeod & Adams, 1968
• Manselliberotha Aspöck & Aspöck, 1988
• †Microberotha Archibald & Makarkin 2004 (Eocene (Ypresian), Hat Creek amber, Canada)[2]
• Ormiscocerus Blanchard, 1851
• †Protoberotha Huang et al., 2019 (Cenomanian, Burmese amber, Myanmar)[6]
• †Sibelliberotha Azar & Nel 2013 (Barremian, Lebanese amber)
• Speleoberotha Machado et al., 2022[5]

Subfamily †Mesithoninae Panfilov, 1980

• †Mesithone Panfilov 1980 Callovian/Oxfordian; Karabastau Formation, Kazakhstan; Aptian; Zaza Formation, Turga Formation, Russia
• †Oloberotha Ren & Guo 1996 Aptian; Yixian Formation, China

Subfamily Nosybinae

• †Banoberotha Whalley 1980 Cretaceous (Barremian); Lebanese amber, Lebanon
• Naizema Navás, 1919
• Nosybus Navás, 1910
• Tanzanberotha Aspöck & Hynd, 1995

Subfamily Nyrminae

• Nyrma Navás, 1933

Subfamily Protobiellinae

• Austroberothella Aspöck & Aspöck, 1985
• Protobiella Tillyard, 1923

Subfamily Trichomatinae

• Trichoberotha Handschin, 1935
• Trichoma Tillyard, 1916

Subfamily Incertae sedis

• †Ansoberotha Yang et al., 2019 Cenomanian; Burmese amber, Myanmar
• †Berothone Khramov 2015 Callovian/Oxfordian; Karabastau Formation, Kazakhstan
• †Cantabroberotha Pérez-de la Fuente et al., 2021 Cretaceous (Albian); Spanish amber, Spain
• †Cornoberotha Yang et al., 2019 Cenomanian; Burmese amber, Myanmar
• †Dasyberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Epimesoberotha Jepson et al., 2012 Cretaceous (Berriasian); Lulworth Formation, United Kingdom
• †Ethiroberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Haploberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Iceloberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Jersiberotha Grimaldi 2000 Cenomanian; Burmese amber, Myanmar; Cretaceous (Turonian); New Jersey amber, United States
• †Krokhathone Khramov 2015 Jurassic (Callovian/Oxfordian); Karabastau Formation, Kazakhstan
• †Maculaberotha Yuan et al., 2016 Cenomanian; Burmese amber, Myanmar
• †Magniberotha Yuan et al., 2016 Cenomanian; Burmese amber, Myanmar
• †Nascimberotha Grimaldi 2000 Turonian; New Jersey amber, United States
• †Plesiorobius Klimaszewski & Kevan, 1986 Cretaceous (Campanian); Canadian amber, Canada; Campanian; Taymyr amber, Russia; Cenomanian, unknown formation, Russia
• †Proberotha Krüger, 1923 Lutetian, Baltic Amber, northern Europe
• †Osmyloberotha Khramov 2021 Cenomanian; Burmese amber, Myanmar
• †Sinosmylites Hong 1983 Daohugou, Haifanggou Formation, China, Callovian, Karabastau Formation, Kazakhstan, Callovian/Oxfordian, Ulaan-Ereg Formation, Mongolia, Late Jurassic (Tithonian)
• †Systenoberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Telistoberotha Engel & Grimaldi 2008 Cenomanian; Burmese amber, Myanmar
• †Xiaoberotha Shi et al., 2019 Cenomanian; Burmese amber, Myanmar

Formerly included taxa

Subfamily Paraberothinae (now included in Rhachiberothidae)

• †Acanthoberotha Nakamine et al., 2020 Cenomanian; Burmese amber, Myanmar
• †Alboberotha Nel et al., 2005 Cretaceous (Albian); Charentense amber, France
• †Astioberotha Nakamine et al., 2020 Cenomanian; Burmese amber, Myanmar
• †Chimerhachiberotha Nel et al., 2005 Barremian; Lebanese amber, Lebanon
• †Creagroparaberotha Makarkin, 2015 Cenomanian; Burmese amber, Myanmar
• †EorhachiberothaEngel, 2004 Cenomanian; Burmese amber, Myanmar
• †Kujiberotha Nakamine & Yamamoto, 2018 Cretaceous (Santonian); Kuji amber, Japan
• †Micromantispa Shi et al., 2015 Cenomanian; Burmese amber, Myanmar
• †Paraberotha Whalley, 1980 Barremian; Lebanese amber, Lebanon
• †Pseudosisyra Makarkin, 1999 Aptian; Zaza Formation, Russia
• †Raptorapax Petrulevicius et al., 2010 Barremian; Lebanese amber, Lebanon
• †Retinoberotha Schlüter, 1978 Cenomanian; Bezonnais amber, France,
• †Rhachibermissa Grimaldi, 2000 Turonian; New Jersey amber, United States
• †Scoloberotha Engel & Grimaldi, 2008 Cenomanian; Burmese amber, Myanmar
• †Spinoberotha Nel et al., 2005 Cenomanian; Burmese amber, Myanmar
• †Stygioberotha Nakamine et al, 2020 Cenomanian; Burmese amber, Myanmar
• †Uranoberotha Nakamine et al, 2020 Cenomanian; Burmese amber, Myanmar

References

1. Winterton, Shaun L. (2010). "A new species of Stenobiella Tillyard (Neuroptera, Berothidae) from Australia". ZooKeys (64): 1–8. doi:10.3897/zookeys.64.403. PMC 3088400. PMID 21594021.
2. Archibald, S.B.; Makarkin, V.N. (2004). "New genus of minute Berothidae (Neuroptera) from Early Eocene amber of British Columbia". The Canadian Entomologist. 136 (1): 61–76. CiteSeerX 10.1.1.552.2285. doi:10.4039/n03-043. S2CID 36459014.
3. Aspöck, Ulrike; Randolf, Susanne (2 December 2014). "Beaded lacewings – a pictorial identification key to the genera, their biogeographics and a phylogentic analysis (Insecta: Neuroptera: Berothidae)". Deutsche Entomologische Zeitschrift. 61 (2): 155–172. doi:10.3897/dez.61.8850 – via Pensoft.
4. Engel, Michael S.; Winterton, Shaun L.; Breitkreuz, Laura C. V. (2018-01-07). "Phylogeny and Evolution of Neuropterida: Where Have Wings of Lace Taken Us?". Annual Review of Entomology. 63: 531–551. doi:10.1146/annurev-ento-020117-043127. ISSN 1545-4487. PMID 29324039.
5. Machado, R. J. P.; Martins, C. C.; Aspöck, H.; De Miranda Tavares, L. G.; Aspöck, U. (2022). "The first cave associated genus of Berothidae (Insecta: Neuroptera), and a new interpretation of the subfamily Cyrenoberothinae". Zoological Journal of the Linnean Society. 195 (4): 1422–1444. doi:10.1093/zoolinnean/zlab104.
6. Huang, S.; Ren, D.; Wang, Y. (2019). "A new basal beaded lacewing (Neuroptera: Berothidae) from mid-Cretaceous Myanmar amber". Cretaceous Research. 95: 1–7. doi:10.1016/j.cretres.2018.10.025.