Owlfly

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.

Owlfly
Temporal range:
both female Libelloides macaronius
Istria, Croatia
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Neuroptera
Clade: Myrmeleontiformia
Family: Ascalaphidae
Rambur, 1842
Subfamilies[1]

and see text

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.[2][3] 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 1.5 inches (3.8 cm) in length, not including antennae.[4] 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.[4] Owlflies are worldwide in distribution, occurring in warm temperate and tropical habitats; there are some 450 extant species.[5]

Ecology
Some owlflies raise the abdomen at rest, mimicking a broken twig.[4]

Adult owlflies are fast-flying, aerial predators, capturing and feeding on other insects in flight.[6] The larvae too are predatory, making owlflies important in maintaining a natural ecological balance and helping to control pest insects.[7]

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.[4] Some Old World species, such as Libelloides macaronius, are active during the day.[8]

Anti-predator defences
Further information: Anti-predator adaptation

When disturbed, some owlflies release a strong, musk-like chemical to deter enemies.[4] The abdomen in Ululodes quadrimaculatus is raised at rest, mimicking a broken twig.[4]

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,[9] or as heliographic signalling, reflecting sunlight, to attract females.[10]

Life cycle
Brood of first instar larvae on their egg-cases before dispersing
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.[11] In some genera, larvae actively place sand and debris onto their dorsum as self-decoration camouflage.[12] Pupation occurs in a spheroidal silk cocoon in leaf litter or soil.[13]

Evolution

Owlflies appear to have evolved from a common ancestor with Stilbopterygidae.[1] These, in turn, evolved from a common ancestor with Palparidae, which evolved from a common ancestor with the true antlions, or Myrmeleontidae.[1]

Taxonomy and etymology

The family Ascalaphidae was first described by the French entomologist Jules Pierre Rambur in 1842.[14] The name is from Greek askalaphos, a kind of owl.[15] In Greek mythology, Ascalaphos was the custodian of the orchard of Hades, god of the underworld; the goddess Demeter transformed him into an owl.[16]

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.[17][18] 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.[19]

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.[1] This refuted nuclear phylogenomic analysis in 2018, which recovered Ascalaphidae as a paraphyletic lineage within Myrmeleontidae.[20] 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.[21][22] The fossil record has contributed to an understanding of the group's phylogeny.[19][23] The phylogeny of the owlflies has remained uncertain, with many of the higher taxa apparently not natural groups (clades).[24]

External

Neuropteran subfamilies are described in Winterton and colleagues 2017 and Jones 2019.[25][1]

Neuropterida

Raphidioptera

Megaloptera

Neuroptera

6 subfamilies[25]

Ithonidae

giant lacewings,
moth lacewings
Myrmeleontiformia
Psychopsidae

silky lacewings
Nymphidae

split‑footed
lacewings
Nemopteridae

spoonwings,
threadwings

Myrmeleontoidea

lacewings

Internal

Machado et al 2018 proposes a classification below family level, into tribes (names ending with –ini):[20] Groups formerly considered part of "Myrmeleontidae" are underscored and marked "Myrm."

Myrmeleontoidea
Ascalaphinae

Dimarini

Palparini "Myrm."

Ululodini

Stilbopterygini "Myrm."

Haplogleniini

Ascalaphini

"Myrmeleontidae" (part)

Myrmeleontinae

Dendroleontinae

Nemoleontinae

Jones 2019 presents a total-evidence phylogeny, preferring to classify only to family level:[1]

Myrmeleontoidea

Myrmeleontidae

Palparidae "Myrm."

Stilbopterygini "Myrm."

Ascalaphidae

Albardiinae

Ululodinae

Melambrotinae

Haplogleniinae

Ascalaphinae

owlflies

References
  1. Jones, Joshua R. (2019). "Total-evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher-level classification". Zoologica Scripta. 48 (6): 761–782. doi:10.1111/zsc.12382.
  2. "Owlflies". Missouri Department of Conservation. Retrieved 26 December 2020.
  3. Foltz, John L. (August 10, 2004). "Neuroptera: Ascalaphidae". ENY 3005 Family Identification. University of Florida. Archived from the original on June 1, 2010. Retrieved July 14, 2010.
  4. Saad, Brooke; Pickens, Lindsey (2004). "Ululodes quadrimaculatus, "owlfly"". Clemson University. Archived from the original on October 12, 2008. Retrieved April 24, 2008.
  5. Trujillo, Gloria (2009). "Neuroptera: Ascalaphidae" (PDF). University of Florida. Retrieved 4 January 2022.
  6. Britton, David (12 March 2019). "Life-history of an Owlfly". Australian Museum.
  7. Gao, Xin-Yan; Cai, Yin-Yin; Yu, Dan-Na; Storey, Kenneth B.; Zhang, Jia-Yong (14 November 2018). "Characteristics of the complete mitochondrial genome of Suhpalacsa longialata (Neuroptera, Ascalaphidae) and its phylogenetic implications". PeerJ. 6: e5914. doi:10.7717/peerj.5914. ISSN 2167-8359. PMC 6240338. PMID 30479895.
  8. Chinery, Michael (1982). A Field Guide to the Insects of Britain and Northern Europe. Collins. p. 151. ISBN 0-002192160.
  9. New, Timothy R. (2014). Kükenthal, Willy (ed.). Planipennia: Lacewings. De Gruyter. p. 51. doi:10.1515/9783110858815. ISBN 978-3-11-085881-5.
  10. Onore, G.; Badano, D.; Pantaleoni, R. A. (2014). "Heliographic signalling in Haploglenius Burmeister, 1839 (Neuroptera Ascalaphidae)" (PDF). Biodiversity Journal. 5 (1): 87–91.
  11. Badano, D.; Pantaleoni, R. A. (2014). "The Larvae of European Ascalaphidae (Neuroptera)" (PDF). Zootaxa. 3796 (2): 287–319. doi:10.11646/zootaxa.3796.2.4. PMID 24870677. S2CID 41165521.
  12. Henry, C. S. (1977). "The behavior and life histories of two North American ascalaphids" (PDF). Annals of the Entomological Society of America. 70 (2): 179–195. doi:10.1093/aesa/70.2.179.
  13. "Family Ascalaphidae - Owlflies". BugGuide. September 19, 2006. Retrieved April 24, 2008.
  14. Rambur, Jules Pierre (1842). Histoire naturelle des insectes, névroptères. Librairie Encyclopédique de Roret. Paris: Fain et Thunot.
  15. "History and Etymology for Ascalaphidae". Merriam-Webster. Retrieved 4 January 2022.
  16. Ovid. Metamorphoses. Vol. V. Brookes More, Boston, Cornhill Publishing Co. p. 534.
  17. MacLeod, Ellis G. (June 1970). "The Neuroptera of the Baltic Amber. 1. Ascalaphidae, Nymphidae, and Psychopsidae" (PDF). Psyche. 77 (2): 147–180. doi:10.1155/1970/45459.
  18. Badano, Davide; Engel, Michael S.; Basso, Andrea; Wang, Bo; Cerretti, Pierfilippo (22 August 2018). "Diverse Cretaceous larvae reveal the evolutionary and behavioural history of antlions and lacewings". Nature Communications. 9 (1). article 3257. Bibcode:2018NatCo...9.3257B. doi:10.1038/s41467-018-05484-y. ISSN 2041-1723. PMC 6105666. PMID 30135436.
  19. Engel, Michael S.; Grimaldi, David A. (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. S2CID 49393365.
  20. Machado, R. J. P.; Gillung, J. P.; Winterton, S. L.; Garzon‐Orduña, I. J.; Lemmon, A. R.; Lemmon, E. M.; Oswald, J. D. (2018). "Owlflies are derived antlions: Anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera)". Systematic Entomology. 44 (2): 418–450. doi:10.1111/syen.12334.
  21. Yue, Bi-Song; Song, Nan; Lin, Aili; Zhao, Xincheng (2018). "Insight into higher-level phylogeny of Neuropterida: Evidence from secondary structures of mitochondrial rRNA genes and mitogenomic data". PLOS ONE. 13 (1): e0191826. Bibcode:2018PLoSO..1391826S. doi:10.1371/journal.pone.0191826. ISSN 1932-6203. PMC 5790268. PMID 29381758.
  22. Yan, Y.; Wang Y, Liu, X.; Winterton, S. L.; Yang, D. (2014). "The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia". International Journal of Biological Sciences. 10 (8): 895–908. doi:10.7150/ijbs.9454. PMC 4147223. PMID 25170303.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  23. Parker, S. P. (ed.), 1982: Synopsis and classification of living organisms. Vols. 1 & 2. McGrew-Hill Book Company
  24. Jones, Joshua Raymond (December 2014). Taxonomic Revisions of Six Genera of Entire-Eyed Owlflies (Ascalaphidae: Haplogleniinae), and First Large-Scale Phylogeny of the Owlflies. Texas A&M University (PhD Thesis).
  25. Winterton, S. L.; Lemmon, A. R.; Gillung, J. P.; Garzon, I. J.; Badano, D.; Bakkes, D. K.; Breitkreuz, L. C. V.; Engel, M. S.; Lemmon, E. M.; Liu, X.; Machado, R. J. P.; Skevington, J. H.; Oswald, J. D. (2017). "Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera)". Systematic Entomology. 43 (2): 330–354. doi:10.1111/syen.12278.
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