Hemerobiidae
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Hemerobiidae is a family of
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
n 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 are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and (differing between sources) 1,300–2,000 species in this widespread group. Members of the genera '' Chrysopa'' and '' Chrysoperl ...
(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 Costal may refer to: * an adjective related to the rib () in anatomy ** Costal cartilage, a type of cartilage forming bars which serve to prolong the ribs forward ** Costal margin, the medial margin formed by the false ribs ** Costal surface (disa ...
cross veins. Some genera (''
Hemerobius ''Hemerobius'' is a genus of lacewings in the family Hemerobiidae. It is found throughout Europe and North America. Like most lacewings, both the larvae and adults are predatory, primarily eating acarines, scale insects, psyllids, aphids, thri ...
'', ''
Micromus ''Micromus'' is a genus of lacewings in the family Hemerobiidae (the brown lacewings). These small (4–10 mm long) insects are found worldwide.The Global Biodiversity Information Facility: GBIF Backbone Taxonomy doi:10.15468/39omei Accesse ...
'', '' Notiobiella'', '' Sympherobius'', ''
Wesmaelius ''Wesmaelius'' is a genus of net-winged insects, belonging to the family Hemerobiidae 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, ...
'') 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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Drepanepteryginae
mimic MIMIC, known in capitalized form only, is a former simulation computer language developed 1964 by H. E. Petersen, F. J. Sansom and L. M. Warshawsky of Systems Engineering Group within the Air Force Materiel Command at the Wright-Patterson AFB in ...
dead leaves. Hemerobiid larvae are usually less hairy than chrysopid larvae. Hemerobiids, like chrysopids, are
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
y, especially on
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s, both as
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e and adults. The
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
''
Micromus tasmaniae ''Micromus tasmaniae'', known as the Tasmanian brown lacewing, is a species of brown lacewing in the family Hemerobiidae. It is widespread in Australia, New Zealand, and Pacific Islands such as New Caledonia and Vanuatu. Description and ecol ...
'' is bred for
biological pest control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
.


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 Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in ...
. The
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Hemerobioidea is currently restricted to the Hemerobiidae. Formerly, the pleasing lacewings (
Dilaridae Dilaridae is a family of Euneuropteran insects in the order Neuroptera, known as "pleasing lacewings". They were formerly placed in the paraphyletic superfamily Hemerobioidea, though the group is currently placed in the monophyletic superfamily ...
), silky lacewings (
Psychopsidae Psychopsidae is a family of winged insects of the order Neuroptera. They are commonly called silky lacewings. The silky lacewings are distinguishable in their adult stage by their spectacularly patterned and pubescent wings, broad wing shape, d ...
), giant lacewings (
Polystoechotidae 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 mo ...
) and as noted above the green lacewings (
Chrysopidae Green lacewings are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and (differing between sources) 1,300–2,000 species in this widespread group. Members of the genera '' Chrysopa'' and '' Chrysoperl ...
) were placed therein too. Of these, only the
Dilaridae Dilaridae is a family of Euneuropteran insects in the order Neuroptera, known as "pleasing lacewings". They were formerly placed in the paraphyletic superfamily Hemerobioidea, though the group is currently placed in the monophyletic superfamily ...
and
Chrysopidae Green lacewings are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and (differing between sources) 1,300–2,000 species in this widespread group. Members of the genera '' Chrysopa'' and '' Chrysoperl ...
seem to be reasonably close relatives of the brown lacewings. The
Psychopsidae Psychopsidae is a family of winged insects of the order Neuroptera. They are commonly called silky lacewings. The silky lacewings are distinguishable in their adult stage by their spectacularly patterned and pubescent wings, broad wing shape, d ...
in fact seem to belong to an altogether different
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
of Neuroptera, the
Myrmeleontiformia Myrmeleontiformia is an insect clade in the order Neuroptera, and which was historically treated as a suborder. The phylogeny of the Neuroptera has been explored using mitochondrial DNA sequences, and while issues remain for the order as a whole ...
.


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 ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''.


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 ''Hemerobius'' is a genus of lacewings in the family Hemerobiidae. It is found throughout Europe and North America. Like most lacewings, both the larvae and adults are predatory, primarily eating acarines, scale insects, psyllids, aphids, thri ...
'' Linnaeus, 1758 ** Genus '' Nesobiella'' ** Genus ''
Wesmaelius ''Wesmaelius'' is a genus of net-winged insects, belonging to the family Hemerobiidae 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, ...
'' Krüger, 1922 * Subfamily Megalominae ** Genus '' Megalomus'' Rambur, 1842 * Subfamily
Microminae Microminae is a subfamily of neuropteran insects of the family Hemerobiidae. Includes the genera '' Micromus'' (with global distribution), '' Nusalala'' (with Neotropical distribution), '' Noius'' (New Caledonia), and '' Megalomina'' (Australia ...
** Genus '' Megalomina'' Banks, 1909 ** Genus ''
Micromus ''Micromus'' is a genus of lacewings in the family Hemerobiidae (the brown lacewings). These small (4–10 mm long) insects are found worldwide.The Global Biodiversity Information Facility: GBIF Backbone Taxonomy doi:10.15468/39omei Accesse ...
'' Rambur, 1842 ** Genus ''
Nusalala ''Nusalala'' is a genus of brown lacewings. The scientific name was published in 1913 by Longinos Navás. They belong to the subfamily Microminae, as well as the genera ''Micromus'' and '' Megalomina''. Some species of this genus, such as ''Nusa ...
'' 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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
, the genus '' Notherobius'' is of uncertain or fairly basal position.


Fossils

Numerous
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
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 The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
to
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
rocks or
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
, '' Promegalomus'' is known from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. It was formerly considered to constitute a distinct family Promegalomidae, but is currently recognized as a very basal member of the Hemerobiidae. The
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
'' Mesohemerobius'' was formerly considered a brown lacewing, but is today rather placed as ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' 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 Quesnel (Kee-nel in French) is a city located in the Cariboo Regional District of British Columbia, Canada. Located nearly evenly between the cities of Prince George and Williams Lake, it is on the main route to northern British Columbia and th ...
) * '' Brasilopsychopsis''
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous ( Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly duri ...
Brazil, Early Cretaceous (
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
) * '' Cratopsychopsis'' Crato Formation Brazil, Aptian * '' Cretomerobius'' Ponomarenko, 1992 Dzun-Bain Formation, Mongolia, Aptian *'' Hemeroberotha'' Makarkin & Gröhn, 2020
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Late Cretaceous (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
) *'' Mucropalpus'' Pictet, 1856 (Eocene;
Baltic Baltic may refer to: Peoples and languages * Baltic languages, a subfamily of Indo-European languages, including Lithuanian, Latvian and extinct Old Prussian *Balts (or Baltic peoples), ethnic groups speaking the Baltic languages and/or originati ...
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
) *'' Plesiorobius'' Klimaszewski and Kevan 1986 Late Cretaceous (
Taimyr amber Taymyr or Taimyr may refer to: Places *Taymyr Peninsula, a peninsula in Siberia *Taymyr Gulf *Taymyra, a river in the Taymyr Peninsula *Lake Taymyr *Taymyr Island, an island in the Kara Sea *Taymyr Autonomous Okrug, a former federal subject of Rus ...
, Russia,
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
Canadian amber Canadians (french: Canadiens) are people identified with the country of Canada. This connection may be residential, legal, historical or cultural. For most Canadians, many (or all) of these connections exist and are collectively the source of ...
,
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...
Ola Formation, Russia, Campanian) * '' Prochlanius'' Kruger, 1923 (Eocene; Baltic amber) * '' Promegalomus'' Panfilov, 1980
Karabastau Formation The Karabastau Formation ( kk, Qarabastaý svıtasy) is a geological formation and lagerstätte in the Karatau Mountains of southern Kazakhstan whose strata date to the Middle to Late Jurassic. It is an important locality for insect fossils that ...
, Kazakhstan, Middle/Late Jurassic (
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
/ Oxfordian) * '' Prophlebonema'' Kruger, 1923 (Eocene; Baltic amber) * '' Prospadobius'' Kruger, 1923 (Eocene; Baltic amber) *'' Purbemerobius'' Jepson et al. 2012
Durlston Formation The Durlston Formation is a geologic formation in England. Particularly in the Isle of Purbeck. It preserves fossils dating back to the Berriasian stage of the Lower Cretaceous. Vertebrate paleobiota Crocodyliformes See also * List of f ...
, United Kingdom, Early Cretaceous (
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...
) 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 Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
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 Qinghai (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest pop ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, in 2018. The finding is from
Lower Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene Epoch made up of two stages: the Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 Ma to 15.97 ± 0.05 Ma (million years ago). It was prece ...
. Some additional brown lacewing larvae have been found as fossils, but it has been impossible to determine their generic or subfamilial association.


References

* (1986): ''Collins Guide to the Insects of Britain and Western Europe''.


External links


BioLib Taxonomic tree of Hemerobiidae (incomplete)
on the UF / IFAS Featured Creatures Web site {{Taxonbar, from=Q790667 Hemerobiiformia Neuroptera Neuroptera families Articles containing video clips Extant Jurassic first appearances