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Hemerobiidae Temporal range: Jurassic–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Adult Micromus variegatus (Microminae)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Neuroptera
Suborder:	Hemerobiiformia
Superfamily:	Hemerobioidea
Family:	Hemerobiidae Latreille, 1802
Subfamilies
10, see text
Synonyms
Promegalomidae

Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long (some up to 18 mm). These insects differ from the somewhat similar Chrysopidae (green lacewings) not only by the usual coloring but also by the wing venation: hemerobiids differ from chrysopids in having numerous long veins (two or more radial sectors) and forked costal cross veins. Some genera (Hemerobius, Micromus, Notiobiella, Sympherobius, Wesmaelius) are widespread, but most are restricted to a single biogeographical realm. Some species have reduced wings to the degree that they are flightless. Imagines (adults) of subfamily Drepanepteryginae mimic dead leaves. Hemerobiid larvae are usually less hairy than chrysopid larvae.

Hemerobiids, like chrysopids, are predatory, especially on aphids, both as larvae and adults. The species Micromus tasmaniae is bred for biological pest control.

Systematics

The relationships between the Hemerobiidae and the other Neuropteran families are still unresolved. Despite their superficial similarity to chrysopids, the brown and green lacewings may not be as closely related as was at one time believed. Rather, the Hemerobiidae appear to be closely related to the Mantispidae.

The superfamily Hemerobioidea is currently restricted to the Hemerobiidae. Formerly, the pleasing lacewings (Dilaridae), silky lacewings (Psychopsidae), giant lacewings (Polystoechotidae) and as noted above the green lacewings (Chrysopidae) were placed therein too. Of these, only the Dilaridae and Chrysopidae seem to be reasonably close relatives of the brown lacewings. The Psychopsidae in fact seem to belong to an altogether different suborder of Neuroptera, the Myrmeleontiformia.

Phylogeny

Cladogram of Hemerobiidae relations, based on morphological and molecular data. Psychobiellinae was rearranged into Notiobiellinae and Zachobiellinae, and Adelphohemerobiinae was placed as incertae sedis.

Genera

The subfamilies of Hemerobiidae are:

• Subfamily Adelphohemerobiinae
  • Genus Adelphohemerobius Oswald, 1993
• Subfamily Drepanacrinae
  • Genus Austromegalomus
  • Genus Conchopterella
  • Genus Drepanacra
• Subfamily Carobiinae
  • Genus Carobius
• Subfamily Drepanepteryginae
  • Genus Drepanepteryx Leach, 1815
  • Genus Gayomyia
  • Genus Neuronema
• Subfamily Hemerobiinae
  • Genus Biramus
  • Genus Hemerobiella
  • Genus Hemerobius Linnaeus, 1758
  • Genus Nesobiella
  • Genus Wesmaelius Krüger, 1922
• Subfamily Megalominae
  • Genus Megalomus Rambur, 1842
• Subfamily Microminae
  • Genus Megalomina Banks, 1909
  • Genus Micromus Rambur, 1842
  • Genus Nusalala Navás, 1913
• Subfamily Notiobiellinae
  • Genus Anapsectra
  • Genus Notiobiella
  • Genus Psectra Hagen, 1866
  • Genus Zachobiella
• Subfamily Psychobiellinae
  • Genus Psychobiella
• Subfamily Sympherobiinae
  • Genus Neosympherobius
  • Genus Nesobiella
  • Genus Sympherobius Banks, 1904

Apart from the genera assigned to subfamilies, the genus Notherobius is of uncertain or fairly basal position.

Fossils

Numerous fossil Hemerobiidae have been described, some from the still-living genera, others from genera that are entirely extinct today. While most have been found in Eocene to Miocene rocks or amber, Promegalomus is known from the Jurassic. It was formerly considered to constitute a distinct family Promegalomidae, but is currently recognized as a very basal member of the Hemerobiidae. The Cretaceous Mesohemerobius was formerly considered a brown lacewing, but is today rather placed as incertae sedis in the Neuroptera; it might be a member of the Hemerobioidea but not even that is certain. Notable fossil Hemerobiidae genera are:

• Bothromicromus Scudder, 1878 (Eocene/Oligocene; Quesnel, British Columbia)
• Brasilopsychopsis Crato Formation Brazil, Early Cretaceous (Aptian)
• Cratopsychopsis Crato Formation Brazil, Aptian
• Cretomerobius Ponomarenko, 1992 Dzun-Bain Formation, Mongolia, Aptian
• Hemeroberotha Makarkin & Gröhn, 2020 Burmese amber, Myanmar, Late Cretaceous (Cenomanian)
• Mucropalpus Pictet, 1856 (Eocene; Baltic amber)
• Plesiorobius Klimaszewski and Kevan 1986 Late Cretaceous (Taimyr amber, Russia, Santonian Canadian amber, Campanian Ola Formation, Russia, Campanian)
• Prochlanius Kruger, 1923 (Eocene; Baltic amber)
• Promegalomus Panfilov, 1980 Karabastau Formation, Kazakhstan, Middle/Late Jurassic (Callovian/Oxfordian)
• Prophlebonema Kruger, 1923 (Eocene; Baltic amber)
• Prospadobius Kruger, 1923 (Eocene; Baltic amber)
• Purbemerobius Jepson et al. 2012 Durlston Formation, United Kingdom, Early Cretaceous (Berriasian)

The extinct genus Hemerobites was originally described in 1813 from a specimen preserved in Baltic amber as a hemerobiid. However further study has resulted in its synonymy with the genus Eutermes and placement in the termite subfamily Termitina. The species Wesmaelius mathewesi, was described in 2003 from a solitary Eocene fossil found near Quesnel, British Columbia, and placed into the extant genus Wesmaelius. At that time it was the oldest Hemerobiinae species described. Another species from the same genus, Wesmaelius makarkini was found in Garang Formation of Zeku County, Qinghai Province, China, in 2018. The finding is from Lower Miocene.

Some additional brown lacewing larvae have been found as fossils, but it has been impossible to determine their generic or subfamilial association.

References

1. Stange, Lionel (2008). "Lacewings, Antlions and Matispids (Neuroptera)". In Capinera, John L. (ed.). Encyclopedia of entomology (2nd. ed.). Dordrecht: Springer. pp. 2102–2110. ISBN 978-1-4020-6242-1.
2. New, TR (2002). "Prospects for extending the use of Australian lacewings in biological control" (PDF). Acta Zoologica Academiae Scientiarum Hungaricae. 48 (Supplement 2): 209–216.
3. ^ Aspoeck, Ulrike; Haring, Elisabeth; Aspoeck, Horst (2012). "The phylogeny of the Neuropterida: long lasting and current controversies and challenges (Insecta: Endopterygota)" (PDF). Arthropod Systematics & Phylogeny. 70 (2): 119–129. doi:10.3897/asp.70.e31758. Retrieved 12 September 2016.
4. Winterton, Shaun L.; Hardy, Nate B.; Wiegmann, Brian M. (July 2010). "On wings of lace: Phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data". Systematic Entomology. 35 (3): 349–378. Bibcode:2010SysEn..35..349W. doi:10.1111/j.1365-3113.2010.00521.x. S2CID 84610713.
5. Zhao, Yang; Chen, Yunjiao; Zhao, Jing; Liu, ZhiQi (2015). "First complete mitochondrial genome from the brown lacewings (Neuroptera: Hemerobiidae)". Mitochondrial DNA Part A. 27 (4): 2763–2764. doi:10.3109/19401736.2015.1053054. PMID 26367792. S2CID 6911900.
6. Garzón-Orduña, IJ; Menchaca-Armenta, I; Contreras-Ramos, A; Liu, X; Winterton, SL (20 September 2016). "The phylogeny of brown lacewings (Neuroptera: Hemerobiidae) reveals multiple reductions in wing venation". BMC Evolutionary Biology. 16 (1): 192. Bibcode:2016BMCEE..16..192G. doi:10.1186/s12862-016-0746-5. PMC 5029026. PMID 27645380.
7. ^ Yang, Qiang; Shi, Chaofan; Li, Xiangchuan; Pang, Hong; Ren, Dong (2018-01-10). "The first fossil brown lacewing from the Miocene of the Tibetan Plateau (Neuroptera, Hemerobiidae)". ZooKeys (726): 145–154. Bibcode:2018ZooK..726..145Y. doi:10.3897/zookeys.726.21086. ISSN 1313-2970. PMC 5806409. PMID 29430206.
8. ^ Engel, MS; Grimaldi, DA (2007). "The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera)" (PDF). American Museum Novitates (3587): 1–58. doi:10.1206/0003-0082(2007)3587[1:tnfoda]2.0.co;2. hdl:2246/5880.
9. Makarkin, VN; Archibald, SB; Oswald, JD (2003). "New Early Eocene brown lacewings (Neuroptera: Hemerobiidae) from western North America". The Canadian Entomologist. 135 (5): 637–653. CiteSeerX 10.1.1.489.5852. doi:10.4039/n02-122. S2CID 53479449.

• Chinery, Michael (1986): Collins Guide to the Insects of Britain and Western Europe.

External links

• Media related to Hemerobiidae at Wikimedia Commons
• BioLib Taxonomic tree of Hemerobiidae (incomplete)
• Brown lacewings of Florida on the UF / IFAS Featured Creatures Web site

• Hemerobiiformia
• Neuroptera
• Neuroptera families
• Extant Jurassic first appearances