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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 Chrysoperla are very common in North America and Europe; they are very similar[1] and many of their species have been moved from one genus to the other time and again, and in the nonscientific literature assignment to Chrysopa and Chrysoperla can rarely be relied upon. Since they are the most familiar neuropterans to many people, they are often simply called "lacewings". Since most of the diversity of Neuroptera are properly referred to as some sort of "lacewing", common lacewings is preferable.

Green lacewings
Temporal range: Late Jurassic–Present
Chrysopa perla
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Neuroptera
Superfamily: Chrysopoidea
Family: Chrysopidae
Subfamilies

Apochrysinae
Chrysopinae
Nothochrysinae
and see text

Description and ecology edit

 
Head close-up of Apertochrysa edwardsi from Austins Ferry, Tasmania, Australia

Green lacewings are delicate insects with a wingspan of 6 to over 65 mm, though the largest forms are tropical. They are characterized by a wide costal field in their wing venation, which includes the cross-veins. The bodies are usually bright green to greenish-brown, and the compound eyes are conspicuously golden in many species. The wings are usually translucent with a slight iridescence; some have green wing veins or a cloudy brownish wing pattern. The vernacular name "stinkflies", used chiefly for Chrysopa species but also for others (e.g. Cunctochrysa) refers to their ability to release a vile smell from paired prothoracic glands when handled.

Adults have tympanal organs at the forewings' base, enabling them to hear well. Some Chrysopa show evasive behavior when they hear a bat's ultrasound calls: when in flight, they close their wings (making their echolocational signature smaller) and drop down to the ground. Green lacewings also use substrate or body vibrations as a form of communication between themselves, especially during courtship. Species which are nearly identical morphologically may sometimes be separated more easily based on their mating signals. For example, the southern European Chrysoperla mediterranea looks almost identical to its northern relative C. carnea (Common Green Lacewing), but their courtship "songs" are very different; individuals of one species will not react to the other's vibrations.[2]

 
Larva of unknown species (from Latvia) camouflaged with sand grains

Adults are crepuscular or nocturnal. They feed on pollen, nectar and honeydew supplemented with mites, aphids and other small arthropods, and some, namely Chrysopa, are mainly predatory. Others feed almost exclusively on nectar and similar substances, and have symbiotic yeasts in their digestive tract to help break down the food into nutrients.[1]

Larvae have either a more slender "humpbacked" shape with a prominent bulge on the thorax, or are plumper, with long bristles jutting out from the sides. These bristles will collect debris and food remains – the empty integuments of aphids, most notably – that provide camouflage from birds.

 
Stalked eggs of unknown species, Mainzer Sand (Rhineland-Palatinate, Germany)
 
Larva of Common Green Lacewing (Chrysoperla carnea) or perhaps C. mediterranea feeding on an aphid

Eggs are deposited at night, singly or in small groups; one female produces some 100–200 eggs. Eggs are placed on plants, usually where aphids are present nearby in numbers. Each egg is hung on a slender stalk about 1 cm long, usually on the underside of a leaf. Immediately after hatching, the larvae moult, then crawls up the egg stalk to feed. They are voracious predators, attacking most insects of suitable size, especially soft-bodied ones (aphids, caterpillars and other insect larvae, insect eggs, and at high population densities also each other). The larvae may also occasionally bite humans, possibly out of either aggression or hunger.[3] Therefore, the larvae are colloquially known as "aphid lions" (also spelled "aphidlions") or "aphid wolves", similar to the related antlions. Their senses are weakly developed, except that they are very sensitive to touch. Walking around in a haphazard fashion, the larvae sway their heads from one side to the other, and when they strike a potential prey object, the larva grasps it. Their maxillae are hollow, allowing a digestive secretion to be injected in the prey; the organs of an aphid can for example be dissolved by this in 90 seconds. Depending on environmental conditions, pupation which takes place in a cocoon takes about 1–3 weeks; species from temperate regions usually overwinter as a prepupa, though C. carnea overwinters as newly hatched adults.

Use in biological pest control edit

While depending on species and environmental conditions, some green lacewings will eat only about 150 prey items in their entire lives, in other cases 100 aphids will be eaten in a single week. Thus, in several countries, millions of such voracious Chrysopidae are reared for sale as biological control agents of insect and mite pests in agriculture and gardens. They are distributed as eggs, since as noted above they are highly aggressive and cannibalistic in confined quarters; the eggs hatch in the field. Their performance is variable; thus, there is interest in further research to improve the use of green lacewings as biological pest control. Species that have hitherto attracted wider study and are more or less readily available as captive-bred eggs to deposit out for hatching in pest-infested plant cultures are several members of Chrysoperla as well as Mallada signatus.[4] They are a natural predator of the European corn borer, a moth that costs the US agriculture industry more than $1 billion annually in crop losses and population control.[5][6]

Gardeners can attract these lacewings – and therefore ensure a steady supply of larvae – by using certain companion plants and tolerating beneficial weeds. Chrysopidae are attracted mainly by Asteraceae – e.g. calliopsis (Coreopsis), cosmos (Cosmos), sunflowers (Helianthus) and dandelion (Taraxacum) – and Apiaceae such as dill (Anethum) or angelica (Angelica).

Systematics and taxonomy edit

 
Chrysopa sp.

For a long time, green lacewings were considered close relatives of the pleasing lacewings (Dilaridae) and brown lacewings (Hemerobiidae) and placed in the superfamily Hemerobioidea. But this grouping does not appear to be natural and misled most significantly by the supposed hemerobioideans' plesiomorphic larvae. Today, the Hemerobioidea are usually considered monotypic, containing only the brown lacewings; the green lacewings seem to be very closely related to the osmylids (Osmylidae), which have much more advanced larvae superficially resembling those of the spongillaflies (Sisyridae) with which the spongillaflies were thus formerly allied. Thus the superfamily Osmyloidea – also monotypic following the spongillaflies' removal from there – is the closest living relative of green lacewings; some Mesozoic taxa have been placed in families even closer to Chrysopidae (Ascalochrysidae and Mesochrysopidae) and united with these to superfamily Chrysopoidea.[7]

Subfamilies and genera edit

 
Paleochrysopa monteilsensis holotype wing
 
Nothochrysa fulviceps (Nothochrysinae)
 
Nothancyla verreauxi (Apochrysinae)

The living genera of Chrysopidae are divided into one very large and two smaller subfamilies; a few genera are not robustly assigned to either of these yet. Compared to other Neuroptera, which have an extensive, sometimes extremely abundant, fossil record, green lacewings are not known from that many fossils, and these are not generally well-studied.[1] Their prehistoric relatives mentioned above, however, indicate that at least the basal radiation of the Chrysopoidea must have happened in the Jurassic already, if not earlier.

Subfamily Apochrysinae Handlirsch, 1908 edit

  • Apochrysa (including Anapochrysa, Lauraya, Nacaura, Oligochrysa, Synthochrysa)[8]
  • Domenechus
  • Joguina – sometimes includes Lainius[8]
  • Lainius Navás, 1913
  • Loyola (including Claverina)[8]
  • Nobilinus
  • Nothancyla

Subfamily Chrysopinae edit

There are over 60 genera placed in four tribes:

  • Ankylopterygini Navas, 1910
    • Six genera
  • Belonopterygini Navas, 1913
    • Fifteen genera
  • Chrysopini Schneider, 1851
    • Thirty-two genera
  • Leucochrysini Adams, 1978
    • Seven genera
  • Incertae sedis
    • Three genera are not placed within any specific tribe

†Subfamily Limaiinae Martins-Neto & Vulcano, 1988 edit

  • Aberrantochrysa Khramov, 2018 (Cretaceous; Russia)
  • Araripechrysa Martins-Neto & Vulcano, 1988 (Cretaceous; Brazil)
  • Baisochrysa Makarkin, 1997 (Jurassic - Cretaceous; Kazakhstan, Russia)
  • Cretachrysa Makrakin, 1994 (Cretaceous; Russia)
  • Drakochrysa Yang & Hong, 1990 (Cretaceous; China)
  • Limaia Martins-Neto & Vulcano, 1988 (Cretaceous; Brazil)
  • Mesypochrysa Martynov, 1927 (Jurassic-Eocene; China, Denmark, Kazakhstan, Mongolia, Russia, United Kingdom)
  • Parabaisochrysa Lu et al., 2018 (Cretaceous; Myanmar)
  • Protochrysa Willmann & Brooks, 1991 (Cretaceous-Eocene; Canada, China, Denmark)

Subfamily Nothochrysinae Navas, 1910 edit

  • Adamsochrysa Makarkin & Archibald, 2013 (Eocene; Canada, United States)
  • Archaeochrysa (Eocene - Oligocene)
  • Asiachrysa Makarkin, 2014 (Eocene; Russia)
  • Asthenochrysa
  • Cimbrochrysa Schlüter, 1982 (Eocene; Denmark, United states)
  • Dictyochrysa
  • Dyspetochrysa (Eocene; United States)
  • Hypochrysa
  • Kimochrysa
  • Leptochrysa Adams & Penny, 1992
  • Lithochrysa (Eocene; Canada, United States)
  • Nothochrysa McLachlan, 1868
  • †Okanaganochrysa Makarkin & Archibald, 2013 (Eocene; Canada)
  • Palaeochrysa Scudder, 1883 (Eocene; United states)
  • Pamochrysa Tjeder, 1966
  • Pimachrysa
  • Pronothochrysa Peñalver et al., 1995 (Miocene, Spain)
  • Pseudochrysopa Makarkin & Archibald, 2013 (Eocene; Canada)
  • Sinonothochrysa Huang et al., 2021 (Paleocene, China)
  • Tribochrysa (Eocene; United States)
  • Triplochrysa

Subfamily Incertae sedis edit

  • Danochrysa Willmann, 1993 (Eocene, Denmark)
  • Sinochrysa Yang, 1992
  • Stephenbrooksia Willmann, 1993 (Eocene, Denmark)
  • Tibetochrysa Yang, 1988

Footnotes edit

  1. ^ a b c Engel & Grimaldi (2007)
  2. ^ Henry et al. (1999)
  3. ^ "Nature's Freddy Krueger". 20 October 2010.
  4. ^ New (2002)
  5. ^ "European corn borer - Ostrinia nubilalis (Hubner)". entnemdept.ufl.edu. Retrieved 2017-11-13.
  6. ^ The European Corn Borer | The European Corn Borer. Retrieved 2017-11-13. {{cite book}}: |website= ignored (help)
  7. ^ See references in Haaramo (2008)
  8. ^ a b c Winterton & Brooks (2002)

References edit

  • Engel, Michael S. & Grimaldi, David A. (2007): The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera). American Museum Novitates 3587: 1–58. PDF fulltext
  • New, T. R. (2002): Prospects for extending the use of Australian lacewings in biological control. Acta Zoologica Academiae Scientiarum Hungaricae 48(Supplement 2): 209–216. PDF fulltext
  • Winterton, S. L. & Brooks, S. J. (2002): Phylogeny of the apochrysine green lacewings (Neuroptera: Chrysopidae: Apochrysinae). Annals of the Entomological Society of America 95(1): 16–28. doi:10.1603/0013-8746(2002)095[0016:POTAGL2.0.CO;2]

Further reading edit

  • Brooks, S. J. & Barnard. P. C. (1990): The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). Bulletin of the British Museum of Natural History (Entomology) 59(2): 117–286.
  • Penny, N. D.; Adams, P. A.; Stange, L. A. (1997): Species Catalog of the Neuroptera, Megaloptera, and Raphidioptera of America North of Mexico. Proceedings of the California Academy of Sciences 50(3): 39–114.
  • Tauber, C. A. (2004): A systematic review of the genus Leucochrysa (Neuroptera: Chrysopidae) in the United States. Annals of the Entomological Society of America 97(6): 1129–1158. doi:10.1603/0013-8746(2004)097[1129:ASROTG2.0.CO;2]  
  • Winterton, S. L. (1995): A new genus and species of Apochrysinae (Neuroptera: Chrysopidae) from Australia, with a checklist of Australian Chrysopidae. Journal of the Australian Entomological Society 34(2): 139–145. doi:10.1111/j.1440-6055.1995.tb01306.x  

External links edit

  •   Media related to Chrysopidae at Wikimedia Commons


chrysopidae, green, lacewings, insects, large, family, order, neuroptera, there, about, genera, differing, between, sources, species, this, widespread, group, members, genera, chrysopa, chrysoperla, very, common, north, america, europe, they, very, similar, ma. 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 Chrysoperla are very common in North America and Europe they are very similar 1 and many of their species have been moved from one genus to the other time and again and in the nonscientific literature assignment to Chrysopa and Chrysoperla can rarely be relied upon Since they are the most familiar neuropterans to many people they are often simply called lacewings Since most of the diversity of Neuroptera are properly referred to as some sort of lacewing common lacewings is preferable Green lacewingsTemporal range Late Jurassic Present PreꞒ Ꞓ O S D C P T J K Pg NChrysopa perlaScientific classificationDomain EukaryotaKingdom AnimaliaPhylum ArthropodaClass InsectaOrder NeuropteraSuperfamily ChrysopoideaFamily ChrysopidaeSubfamiliesApochrysinaeChrysopinaeNothochrysinae and see text Contents 1 Description and ecology 1 1 Use in biological pest control 2 Systematics and taxonomy 3 Subfamilies and genera 3 1 Subfamily Apochrysinae Handlirsch 1908 3 2 Subfamily Chrysopinae 3 3 Subfamily Limaiinae Martins Neto amp Vulcano 1988 3 4 Subfamily Nothochrysinae Navas 1910 3 5 Subfamily Incertae sedis 4 Footnotes 5 References 6 Further reading 7 External linksDescription and ecology edit nbsp Head close up of Apertochrysa edwardsi from Austins Ferry Tasmania AustraliaGreen lacewings are delicate insects with a wingspan of 6 to over 65 mm though the largest forms are tropical They are characterized by a wide costal field in their wing venation which includes the cross veins The bodies are usually bright green to greenish brown and the compound eyes are conspicuously golden in many species The wings are usually translucent with a slight iridescence some have green wing veins or a cloudy brownish wing pattern The vernacular name stinkflies used chiefly for Chrysopa species but also for others e g Cunctochrysa refers to their ability to release a vile smell from paired prothoracic glands when handled Adults have tympanal organs at the forewings base enabling them to hear well Some Chrysopa show evasive behavior when they hear a bat s ultrasound calls when in flight they close their wings making their echolocational signature smaller and drop down to the ground Green lacewings also use substrate or body vibrations as a form of communication between themselves especially during courtship Species which are nearly identical morphologically may sometimes be separated more easily based on their mating signals For example the southern European Chrysoperla mediterranea looks almost identical to its northern relative C carnea Common Green Lacewing but their courtship songs are very different individuals of one species will not react to the other s vibrations 2 nbsp Larva of unknown species from Latvia camouflaged with sand grainsAdults are crepuscular or nocturnal They feed on pollen nectar and honeydew supplemented with mites aphids and other small arthropods and some namely Chrysopa are mainly predatory Others feed almost exclusively on nectar and similar substances and have symbiotic yeasts in their digestive tract to help break down the food into nutrients 1 Larvae have either a more slender humpbacked shape with a prominent bulge on the thorax or are plumper with long bristles jutting out from the sides These bristles will collect debris and food remains the empty integuments of aphids most notably that provide camouflage from birds nbsp Stalked eggs of unknown species Mainzer Sand Rhineland Palatinate Germany nbsp Larva of Common Green Lacewing Chrysoperla carnea or perhaps C mediterranea feeding on an aphidEggs are deposited at night singly or in small groups one female produces some 100 200 eggs Eggs are placed on plants usually where aphids are present nearby in numbers Each egg is hung on a slender stalk about 1 cm long usually on the underside of a leaf Immediately after hatching the larvae moult then crawls up the egg stalk to feed They are voracious predators attacking most insects of suitable size especially soft bodied ones aphids caterpillars and other insect larvae insect eggs and at high population densities also each other The larvae may also occasionally bite humans possibly out of either aggression or hunger 3 Therefore the larvae are colloquially known as aphid lions also spelled aphidlions or aphid wolves similar to the related antlions Their senses are weakly developed except that they are very sensitive to touch Walking around in a haphazard fashion the larvae sway their heads from one side to the other and when they strike a potential prey object the larva grasps it Their maxillae are hollow allowing a digestive secretion to be injected in the prey the organs of an aphid can for example be dissolved by this in 90 seconds Depending on environmental conditions pupation which takes place in a cocoon takes about 1 3 weeks species from temperate regions usually overwinter as a prepupa though C carnea overwinters as newly hatched adults Use in biological pest control edit While depending on species and environmental conditions some green lacewings will eat only about 150 prey items in their entire lives in other cases 100 aphids will be eaten in a single week Thus in several countries millions of such voracious Chrysopidae are reared for sale as biological control agents of insect and mite pests in agriculture and gardens They are distributed as eggs since as noted above they are highly aggressive and cannibalistic in confined quarters the eggs hatch in the field Their performance is variable thus there is interest in further research to improve the use of green lacewings as biological pest control Species that have hitherto attracted wider study and are more or less readily available as captive bred eggs to deposit out for hatching in pest infested plant cultures are several members of Chrysoperla as well as Mallada signatus 4 They are a natural predator of the European corn borer a moth that costs the US agriculture industry more than 1 billion annually in crop losses and population control 5 6 Gardeners can attract these lacewings and therefore ensure a steady supply of larvae by using certain companion plants and tolerating beneficial weeds Chrysopidae are attracted mainly by Asteraceae e g calliopsis Coreopsis cosmos Cosmos sunflowers Helianthus and dandelion Taraxacum and Apiaceae such as dill Anethum or angelica Angelica Systematics and taxonomy edit nbsp Chrysopa sp For a long time green lacewings were considered close relatives of the pleasing lacewings Dilaridae and brown lacewings Hemerobiidae and placed in the superfamily Hemerobioidea But this grouping does not appear to be natural and misled most significantly by the supposed hemerobioideans plesiomorphic larvae Today the Hemerobioidea are usually considered monotypic containing only the brown lacewings the green lacewings seem to be very closely related to the osmylids Osmylidae which have much more advanced larvae superficially resembling those of the spongillaflies Sisyridae with which the spongillaflies were thus formerly allied Thus the superfamily Osmyloidea also monotypic following the spongillaflies removal from there is the closest living relative of green lacewings some Mesozoic taxa have been placed in families even closer to Chrysopidae Ascalochrysidae and Mesochrysopidae and united with these to superfamily Chrysopoidea 7 Subfamilies and genera edit nbsp Paleochrysopa monteilsensis holotype wing nbsp Nothochrysa fulviceps Nothochrysinae nbsp Nothancyla verreauxi Apochrysinae The living genera of Chrysopidae are divided into one very large and two smaller subfamilies a few genera are not robustly assigned to either of these yet Compared to other Neuroptera which have an extensive sometimes extremely abundant fossil record green lacewings are not known from that many fossils and these are not generally well studied 1 Their prehistoric relatives mentioned above however indicate that at least the basal radiation of the Chrysopoidea must have happened in the Jurassic already if not earlier Subfamily Apochrysinae Handlirsch 1908 edit Apochrysa including Anapochrysa Lauraya Nacaura Oligochrysa Synthochrysa 8 Domenechus Joguina sometimes includes Lainius 8 Lainius Navas 1913 Loyola including Claverina 8 Nobilinus Nothancyla Subfamily Chrysopinae edit Main article Chrysopinae There are over 60 genera placed in four tribes Ankylopterygini Navas 1910 Six genera Belonopterygini Navas 1913 Fifteen genera Chrysopini Schneider 1851 Thirty two genera Leucochrysini Adams 1978 Seven genera Incertae sedis Three genera are not placed within any specific tribe Subfamily Limaiinae Martins Neto amp Vulcano 1988 edit Aberrantochrysa Khramov 2018 Cretaceous Russia Araripechrysa Martins Neto amp Vulcano 1988 Cretaceous Brazil Baisochrysa Makarkin 1997 Jurassic Cretaceous Kazakhstan Russia Cretachrysa Makrakin 1994 Cretaceous Russia Drakochrysa Yang amp Hong 1990 Cretaceous China Limaia Martins Neto amp Vulcano 1988 Cretaceous Brazil Mesypochrysa Martynov 1927 Jurassic Eocene China Denmark Kazakhstan Mongolia Russia United Kingdom Parabaisochrysa Lu et al 2018 Cretaceous Myanmar Protochrysa Willmann amp Brooks 1991 Cretaceous Eocene Canada China Denmark Subfamily Nothochrysinae Navas 1910 edit Adamsochrysa Makarkin amp Archibald 2013 Eocene Canada United States Archaeochrysa Eocene Oligocene Asiachrysa Makarkin 2014 Eocene Russia Asthenochrysa Cimbrochrysa Schluter 1982 Eocene Denmark United states Dictyochrysa Dyspetochrysa Eocene United States Hypochrysa Kimochrysa Leptochrysa Adams amp Penny 1992 Lithochrysa Eocene Canada United States Nothochrysa McLachlan 1868 Okanaganochrysa Makarkin amp Archibald 2013 Eocene Canada Palaeochrysa Scudder 1883 Eocene United states Pamochrysa Tjeder 1966 Pimachrysa Pronothochrysa Penalver et al 1995 Miocene Spain Pseudochrysopa Makarkin amp Archibald 2013 Eocene Canada Sinonothochrysa Huang et al 2021 Paleocene China Tribochrysa Eocene United States Triplochrysa Subfamily Incertae sedis edit Danochrysa Willmann 1993 Eocene Denmark Sinochrysa Yang 1992 Stephenbrooksia Willmann 1993 Eocene Denmark Tibetochrysa Yang 1988Footnotes edit a b c Engel amp Grimaldi 2007 Henry et al 1999 Nature s Freddy Krueger 20 October 2010 New 2002 European corn borer Ostrinia nubilalis Hubner entnemdept ufl edu Retrieved 2017 11 13 The European Corn Borer The European Corn Borer Retrieved 2017 11 13 a href Template Cite book html title Template Cite book cite book a website ignored help See references in Haaramo 2008 a b c Winterton amp Brooks 2002 References editEngel Michael S amp Grimaldi David A 2007 The neuropterid fauna of Dominican and Mexican amber Neuropterida Megaloptera Neuroptera American Museum Novitates 3587 1 58 PDF fulltext New T R 2002 Prospects for extending the use of Australian lacewings in biological control Acta Zoologica Academiae Scientiarum Hungaricae 48 Supplement 2 209 216 PDF fulltext Winterton S L amp Brooks S J 2002 Phylogeny of the apochrysine green lacewings Neuroptera Chrysopidae Apochrysinae Annals of the Entomological Society of America 95 1 16 28 doi 10 1603 0013 8746 2002 095 0016 POTAGL2 0 CO 2 PDF fulltextFurther reading editBrooks S J amp Barnard P C 1990 The green lacewings of the world a generic review Neuroptera Chrysopidae Bulletin of the British Museum of Natural History Entomology 59 2 117 286 Penny N D Adams P A Stange L A 1997 Species Catalog of the Neuroptera Megaloptera and Raphidioptera of America North of Mexico Proceedings of the California Academy of Sciences 50 3 39 114 Tauber C A 2004 A systematic review of the genus Leucochrysa Neuroptera Chrysopidae in the United States Annals of the Entomological Society of America 97 6 1129 1158 doi 10 1603 0013 8746 2004 097 1129 ASROTG2 0 CO 2 nbsp Winterton S L 1995 A new genus and species of Apochrysinae Neuroptera Chrysopidae from Australia with a checklist of Australian Chrysopidae Journal of the Australian Entomological Society 34 2 139 145 doi 10 1111 j 1440 6055 1995 tb01306 x nbsp External links edit nbsp Media related to Chrysopidae at Wikimedia Commons Retrieved from https en wikipedia org w index php title Chrysopidae amp oldid 1188872533, wikipedia, wiki, book, books, library,

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