Berothidae

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]

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

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