Owlfly

{{Short description|Family of insects}} {{good article}} {{Automatic taxobox | fossil_range = {{fossil_range|Eocene|Recent}} | image = Owlfly (Libelloides macaronius) female Istria.jpg | image2 = Owlfly (Libelloides macaronius) female Istria 2.jpg | image2_caption = both female ''Libelloides macaronius'' Istria, Croatia | taxon = Ascalaphidae | authority = Rambur, 1842 | subdivision_ranks = Subfamilies | subdivision = * Albardiinae * Ascalaphinae * Haplogleniinae * Melambrotinae * Ululodinae and see text | subdivision_ref = <ref name="Jones2019" /> }}

'''Ascalaphidae''' is a family of insects in the order Neuroptera, commonly called '''owlflies'''; there are some 450 extant species. They are fast-flying crepuscular or diurnal predators of other flying insects, and have large bulging eyes and strongly knobbed antennae. The larvae are ambush predators; some of them make use of self-decoration camouflage.

==Description==

Owlflies are readily distinguished from the superficially similar dragonflies by their long, clubbed antennae; dragonflies have short, bristle-like antennae. The closely related antlions (family Myrmeleontidae) have short, weakly clubbed antennae, smaller eyes, and reticulate wing venation.<ref>{{cite web |title=Owlflies |url=https://nature.mdc.mo.gov/discover-nature/field-guide/owlflies |publisher=Missouri Department of Conservation |access-date=26 December 2020}}</ref><ref>{{cite web |author=Foltz, John L. |work=ENY 3005 Family Identification |url=http://entomology.ifas.ufl.edu/foltz/eny3005/lab1/neuroptera/ascalaphid.htm |title=Neuroptera: Ascalaphidae |date=August 10, 2004 |access-date=July 14, 2010 |publisher=University of Florida |archive-url=https://web.archive.org/web/20100601024018/http://entomology.ifas.ufl.edu/foltz/eny3005/lab1/Neuroptera/ascalaphid.htm |archive-date=June 1, 2010 }}</ref> All but one species of Ascalaphidae have long antennae, easily distinguishing them. The sole exception is the Brazilian ''Albardia furcata'', the only living member of the subfamily Albardiinae, which has short antennae, but these are strongly clubbed (compared to myrmeleontids), and its wing venation is reticulate, typical of ascalaphids. Most owlflies are about {{convert|1.5|in|cm}} in length, not including antennae.<ref name="Clemson"/> Adult owlflies of the family Ululodinae such as ''Ululodes'' have large divided eyes and crepuscular habits, which is where the common name "owlfly" came from.<ref name="Clemson"/> Owlflies are worldwide in distribution, occurring in warm temperate and tropical habitats; there are some 450 extant species.<ref>{{cite web |last1=Trujillo |first1=Gloria |title=Neuroptera: Ascalaphidae |url=https://entnemdept.ufl.edu/choate/neuroptera_ascalaphidae1.pdf |publisher=University of Florida |access-date=4 January 2022 |date=2009 |archive-date=18 April 2024 |archive-url=https://web.archive.org/web/20240418095938/https://entnemdept.ufl.edu/choate/neuroptera_ascalaphidae1.pdf |url-status=dead }}</ref>

==Ecology== [[File:Owlfly.JPG|thumb|upright|Some owlflies raise the abdomen at rest, mimicking a broken twig.<ref name="Clemson"/>]]

Adult owlflies are fast-flying, aerial predators, capturing and feeding on other insects in flight.<ref name="Australian Museum">{{cite web |last=Britton |first=David |title=Life-history of an Owlfly |url=https://australianmuseum.net.au/learn/animals/insects/life-history-of-an-owlfly/ |website=Australian Museum |date=12 March 2019}}</ref> The larvae too are predatory, making owlflies important in maintaining a natural ecological balance and helping to control pest insects.<ref name="Gao Cai Yu Storey p=e5914">{{cite journal |last1=Gao |first1=Xin-Yan |last2=Cai |first2=Yin-Yin |last3=Yu |first3=Dan-Na |last4=Storey |first4=Kenneth B. |last5=Zhang |first5=Jia-Yong |title=Characteristics of the complete mitochondrial genome of Suhpalacsa longialata (Neuroptera, Ascalaphidae) and its phylogenetic implications |journal=PeerJ |volume=6 |date=14 November 2018 |issn=2167-8359 |doi=10.7717/peerj.5914 |article-number=e5914|pmid=30479895 |pmc=6240338 |doi-access=free }}</ref>

Adults of many New World species are most active at sunset, and can often be collected near lights. During the day, adults rest on stems and twigs with the body, legs, and antennae typically pressed to the stem.<ref name="Clemson">{{cite web |last1=Saad |first1=Brooke |last2=Pickens |first2=Lindsey |year=2004 |url=http://entweb.clemson.edu/museum/webonly/local/lmisc/lmisc42.htm |title=''Ululodes quadrimaculatus'', "owlfly" |access-date=April 24, 2008 |publisher=Clemson University |archive-url=https://web.archive.org/web/20081012001128/http://entweb.clemson.edu/museum/webonly/local/lmisc/lmisc42.htm |archive-date=October 12, 2008 }}</ref> Some Old World species, such as ''Libelloides macaronius'', are active during the day.<ref>{{cite book |last=Chinery |first=Michael |author-link=Michael Chinery |title=A Field Guide to the Insects of Britain and Northern Europe |date=1982 |publisher=Collins |isbn=0-002192160 |page=151}}</ref>

===Anti-predator defences=== {{Further|Anti-predator adaptation}}

When disturbed, some owlflies release a strong, musk-like chemical to deter enemies.<ref name="Clemson"/> The abdomen in ''Ululodes quadrimaculatus'' is raised at rest, mimicking a broken twig.<ref name="Clemson"/>

Some New World species such as ''Haploglenius luteus'' are able to suddenly reflex a flap on the pronotum, exposing a strongly-contrasting patch of pale colour (white or cream), either as a deimatic display to startle predators,<ref>{{cite book |last=New |first=Timothy R. |editor-first1=Willy |editor-last1=Kükenthal |title=Planipennia: Lacewings |url=https://books.google.com/books?id=IU1bDAAAQBAJ&pg=PA51 |year=2014 |publisher=De Gruyter |isbn=978-3-11-085881-5 |page=51|doi=10.1515/9783110858815}}</ref> or as heliographic signalling, reflecting sunlight, to attract females.<ref name="Onore2014">{{cite journal |last1= Onore |first1= G. |last2= Badano |first2= D. |last3= Pantaleoni |first3= R. A. |title= Heliographic signalling in Haploglenius Burmeister, 1839 (Neuroptera Ascalaphidae) |journal= Biodiversity Journal |volume=5 |issue=1 |year=2014 |pages=87–91 |url=http://www.biodiversityjournal.com/pdf/5(1)_87-91.pdf }}</ref>

==Life cycle== [[File:Ascalaphidae brood Marita Beneke.jpg|thumb|upright=1.5|Brood of first instar larvae on their egg-cases before dispersing]] thumb|Larva

Eggs are laid on twigs or plant stems. Owlfly larvae are ambush predators, and sequester themselves at the soil surface, in ground litter, or on vegetation, sometimes covered with debris, and wait for prey, which they seize with their large, toothed mandibles. They resemble antlion larvae, but have an elongate, sometimes finger-like appendage on the side of each segment called a scolus-like process.<ref name="Badano2014">{{cite journal | last1= Badano | first1= D. | last2= Pantaleoni | first2= R. A. | title=The Larvae of European Ascalaphidae (Neuroptera) | journal= Zootaxa | volume=3796 | year=2014 |issue=2 | pages=287–319 | doi=10.11646/zootaxa.3796.2.4| pmid= 24870677 | s2cid= 41165521 | url= http://puma.isti.cnr.it/rmydownload.php?filename=cnr.ise/cnr.ise/2014-A0-054/2014-A0-054.pdf }}</ref> In some genera, larvae actively place sand and debris onto their dorsum as self-decoration camouflage.<ref>{{cite journal |last=Henry |first=C. S. |year=1977 |title=The behavior and life histories of two North American ascalaphids |journal=Annals of the Entomological Society of America |volume=70 |issue=2 |pages=179–195 |doi=10.1093/aesa/70.2.179 |url=http://lacewing.tamu.edu/neuropterida/neur_bibliography/edoc12/henry1977ref2877s-4986.pdf}}</ref> Pupation occurs in a spheroidal silk cocoon in leaf litter or soil.<ref>{{cite web |url=http://bugguide.net/node/view/4129 |title=Family Ascalaphidae - Owlflies |date=September 19, 2006 |access-date=April 24, 2008 |publisher=BugGuide}}</ref>

==Evolution== Owlflies appear to have evolved from a common ancestor with Stilbopterygidae.<ref name="Jones2019">{{cite journal | last=Jones | first= Joshua R. | title=Total-evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher-level classification | journal=Zoologica Scripta | volume=48 | year=2019 | issue= 6 | pages=761–782 | doi=10.1111/zsc.12382| doi-access=free }}</ref> These, in turn, evolved from a common ancestor with Palparidae, which evolved from a common ancestor with the true antlions, or Myrmeleontidae.<ref name="Jones2019" />

===Taxonomy and etymology===

The family Ascalaphidae was first described by the French entomologist Jules Pierre Rambur in 1842.<ref>{{cite book |last=Rambur |first=Jules Pierre |author-link=Jules Pierre Rambur |title=Histoire naturelle des insectes, névroptères |series=Librairie Encyclopédique de Roret |publisher=Fain et Thunot |location=Paris |year=1842}}</ref> The name is from Greek ''askalaphos'', a kind of owl.<ref>{{cite web |title=History and Etymology for Ascalaphidae |url=https://www.merriam-webster.com/dictionary/Ascalaphidae |publisher=Merriam-Webster |access-date=4 January 2022}}</ref> In Greek mythology, Ascalaphos was the custodian of the orchard of Hades, god of the underworld; the goddess Demeter transformed him into an owl.<ref>{{cite book |last=Ovid |title=Metamorphoses |volume=V |publisher=Brookes More, Boston, Cornhill Publishing Co. |page=534 |url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.02.0028%3Abook%3D1%3Acard%3D1}}</ref>

===Fossil history===

The owlflies are known from fossils of adults and larvae, often encased in Baltic amber. Most of these cannot be placed in a particular subfamily. Most are known from the Oligocene.<ref>{{cite journal |last1=MacLeod |first1=Ellis G. |title=The Neuroptera of the Baltic Amber. 1. Ascalaphidae, Nymphidae, and Psychopsidae |journal=Psyche: A Journal of Entomology |date=June 1970 |volume=77 |issue=2 |pages=147–180 |url=https://downloads.hindawi.com/journals/psyche/1970/045459.pdf|doi=10.1155/1970/45459|doi-access=free}}</ref><ref name="Badano Engel Basso Wang 2018">{{cite journal |last1=Badano |first1=Davide |last2=Engel |first2=Michael S. |last3=Basso |first3=Andrea |last4=Wang |first4=Bo |last5=Cerretti |first5=Pierfilippo |title=Diverse Cretaceous larvae reveal the evolutionary and behavioural history of antlions and lacewings |journal=Nature Communications |volume=9 |issue=1 |date=22 August 2018 |issn=2041-1723 |doi=10.1038/s41467-018-05484-y |pmid=30135436 |pmc=6105666 |bibcode=2018NatCo...9.3257B |at=article 3257}}</ref> The Late Jurassic ''Mesascalaphus'' was thought to be a more basal member of the family, but it is now believed to be a member of Mesochrysopidae.<ref name="Engel">{{cite journal |author1=Engel, Michael S. |author2=Grimaldi, David A. |year=2007 |title=The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera) |journal=American Museum Novitates |issue=3587 |pages=1–58 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/5880/1/N3587.pdf |doi=10.1206/0003-0082(2007)3587[1:TNFODA]2.0.CO;2|s2cid=49393365 }}</ref>

===Phylogeny===

Total evidence analysis (several genes + morphology) in 2019 recovered Ascalaphidae as monophyletic and found evidence for five subfamilies: Albardiinae van der Weele, 1909; Ululodinae van der Weele, 1909; Haplogleniinae Newman, 1853; Melambrotinae Tjeder, 1992; and Ascalaphinae Lefèbvre, 1842.<ref name="Jones2019" /> This refuted nuclear phylogenomic analysis in 2018, which recovered Ascalaphidae as a paraphyletic lineage within Myrmeleontidae.<ref name="Machado2018">{{cite journal | last1=Machado | first1=R. J. P. | last2=Gillung | first2=J. P. | last3=Winterton | first3=S. L. | last4=Garzon-Orduña | first4=I. J. | last5=Lemmon | first5=A. R. | last6=Lemmon | first6=E. M. | last7=Oswald | first7=J. D. | title=Owlflies are derived antlions: Anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera) | journal=Systematic Entomology | volume=44 | year=2018 | issue=2 | pages=418–450 | doi=10.1111/syen.12334| doi-access=free }}</ref> Molecular analysis in 2018 using mitochondrial rRNA and mitogenomic data also placed the Ascalaphidae as sister to the Myrmeleontidae as the most advanced groups within the Neuroptera.<ref name="YueSong2018">{{cite journal |last1=Yue |first1=Bi-Song |last2=Song |first2=Nan |last3=Lin |first3=Aili |last4=Zhao |first4=Xincheng |title=Insight into higher-level phylogeny of Neuropterida: Evidence from secondary structures of mitochondrial rRNA genes and mitogenomic data |journal=PLOS ONE |volume=13 |issue=1 |year=2018 |article-number=e0191826 |issn=1932-6203 |doi=10.1371/journal.pone.0191826|pmid=29381758 |pmc=5790268 |bibcode=2018PLoSO..1391826S |doi-access=free }}</ref><ref name=Yan>{{cite journal |author1=Yan, Y. |author2=Wang Y, Liu, X. |author3=Winterton, S. L. |author4=Yang, D. |title=The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia |journal=International Journal of Biological Sciences |date=2014 |volume=10 |issue=8 |pages=895–908 |doi=10.7150/ijbs.9454 |pmid=25170303 |pmc=4147223 }}</ref> The fossil record has contributed to an understanding of the group's phylogeny.<ref name="Engel" /><ref>Parker, S. P. (ed.), 1982: Synopsis and classification of living organisms. Vols. 1 & 2. McGrew-Hill Book Company</ref> The phylogeny of the owlflies has remained uncertain, with many of the higher taxa apparently not natural groups (clades).<ref name="Jones 2014">{{cite book |last=Jones |first=Joshua Raymond |title=Taxonomic Revisions of Six Genera of Entire-Eyed Owlflies (Ascalaphidae: Haplogleniinae), and First Large-Scale Phylogeny of the Owlflies |date=December 2014 |publisher=Texas A&M University (PhD Thesis) |url=https://core.ac.uk/reader/147242600}}</ref>

==== External ====

Neuropteran subfamilies are described in Winterton and colleagues 2017 and Jones 2019.<ref name="Winterton2017">{{cite journal | last1=Winterton | first1=S. L. | last2=Lemmon | first2=A. R. | last3=Gillung | first3=J. P. | last4=Garzon | first4=I. J. | last5=Badano | first5=D. | last6=Bakkes | first6=D. K. | last7=Breitkreuz | first7=L. C. V. | last8=Engel | first8=M. S. | last9=Lemmon | first9=E. M. | last10=Liu | first10=X. | last11=Machado | first11=R. J. P. | last12=Skevington | first12=J. H. | last13=Oswald | first13=J. D. | title=Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera) | journal=Systematic Entomology | volume=43 | year=2017 | issue=2 | pages=330–354 | doi=10.1111/syen.12278| doi-access=free | hdl=11573/1554644 | hdl-access=free }}</ref><ref name="Jones2019"/>

{{clade |label1=Neuropterida |1={{clade |1=Raphidioptera |2={{clade |1=Megaloptera |label2=Neuroptera |sublabel2=lacewings |2={{clade |1=6 subfamilies<ref name="Winterton2017"/> |2={{clade |label1=Ithonidae |sublabel1=giant lacewings, moth lacewings |1=75px |label2=Myrmeleontiformia |2={{clade |label1=Psychopsidae |sublabel1=silky lacewings |1=75px |2={{clade |label1=Nymphidae |sublabel1=split-footed lacewings |1=75px |2={{clade |label1=Nemopteridae |sublabel1=spoonwings, threadwings |1=75px |2=Myrmeleontoidea }} }} }} }} }} }} }} }}

==== Internal ====

Machado et al 2018 proposes a classification below family level, into tribes (names ending with –ini):<ref name="Machado2018"/> Groups formerly considered part of "Myrmeleontidae" are underscored and marked "Myrm."

{{clade |label1=Myrmeleontoidea |1={{clade |label1='''Ascalaphinae'''  |1={{clade |1=Dimarini 70px |2={{clade |1=Palparini "Myrm." 75px |2={{clade |1=Ululodini 75px |2={{clade |1=Stilbopterygini "Myrm." 75px |2={{clade |1=Haplogleniini 75px |2=Ascalaphini 75px }} }} }} }} }} |label2="Myrmeleontidae" (part) |2={{clade |1=Myrmeleontinae 100px |2={{clade |1=Dendroleontinae 50px |2=Nemoleontinae 65px }} }} }} }}

Jones 2019 presents a total-evidence phylogeny, preferring to classify only to family level:<ref name="Jones2019"/>

{{clade |label1=Myrmeleontoidea |1={{clade |1=Myrmeleontidae 100px |2={{clade |1=Palparidae "Myrm." 75px |2={{clade |1=Stilbopterygini "Myrm." 75px |2={{clade |label1='''Ascalaphidae''' |sublabel1=owlflies |1={{clade |1=Albardiinae |2={{clade |1=Ululodinae 75px |2={{clade |1=Melambrotinae |2={{clade |1=Haplogleniinae 75px |2=Ascalaphinae 75px }} }} }} }} }} }} }} }} }}

== References == {{Reflist|28em}}

== External links == * {{Wikispecies-inline|Ascalaphidae}} * [https://web.archive.org/web/20131203051457/http://www.asahi-net.or.jp/~CH2M-NITU/tunote.htm Owlflies on Stamps]

Category:Ascalaphidae Category:Neuroptera Category:Neuroptera families Category:Extant Oligocene first appearances Category:Taxa named by Jules Pierre Rambur