Owlfly
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Ascalaphidae is a family of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s in the order Neuroptera, commonly called owlflies; there are some 450 extant species. They are fast-flying
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
or diurnal predators of other flying insects, and have large bulging eyes and strongly knobbed antennae. The larvae are
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey us ...
s; some of them make use of self-decoration camouflage.


Description

Owlflies are readily distinguished from the superficially similar
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
by their long, clubbed antennae; dragonflies have short, bristle-like antennae. The closely related antlions (family
Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the larvae ...
) have short, weakly clubbed antennae, smaller eyes, and reticulate
wing venation Insect wings are adult outgrowths of the insect exoskeleton that enable insects to flight, fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often referre ...
. 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 in length, not including antennae. Adult owlflies of the family
Ululodinae Ululodinae is a subfamily or tribe (as Ululodini) of owlflies. Both they and the Ascalaphinae are sometimes known as split-eyed owlflies due to the characteristic ridge that bisects their compound eyes A compound eye is a visual organ foun ...
such as ''
Ululodes ''Ululodes'' is a genus of owlflies 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, an ...
'' have large divided eyes and
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
habits, which is where the common name "owlfly" came from. Owlflies are worldwide in distribution, occurring in warm temperate and tropical habitats; there are some 450 extant species.


Ecology

Adult owlflies are fast-flying, aerial
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 ...
s, capturing and feeding on other insects in flight. The larvae too are predatory, making owlflies important in maintaining a natural ecological balance and helping to control pest insects. 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. Some Old World species, such as ''
Libelloides macaronius ''Libelloides macaronius'' is a day-flying owlfly species of Europe and Asia. The genus belongs to the family Ascalaphidae, subfamily Ascalaphinae. The species has appeared on postage stamps of Moldova and Ukraine. Description The adult is a ...
'', are active during the day.


Anti-predator defences

When disturbed, some owlflies release a strong,
musk Musk (Persian: مشک, ''Mushk'') is a class of aromatic substances commonly used as base notes in perfumery. They include glandular secretions from animals such as the musk deer, numerous plants emitting similar fragrances, and artificial sub ...
-like chemical to deter enemies. The
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
in ''Ululodes quadrimaculatus'' is raised at rest, mimicking a broken twig. Some New World species such as ''Haploglenius luteus'' are able to suddenly reflex a flap on the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
, exposing a strongly-contrasting patch of pale colour (white or cream), either as a
deimatic display Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anim ...
to startle predators, or as heliographic signalling, reflecting sunlight, to attract females.


Life cycle

Eggs are laid on twigs or plant stems. Owlfly
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 are
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey us ...
s, 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 The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the larvae ...
larvae, but have an elongate, sometimes finger-like appendage on the side of each segment called a scolus-like process. In some genera, larvae actively place sand and debris onto their dorsum as self-decoration camouflage.
Pupation A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
occurs in a spheroidal
silk cocoon A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
in leaf litter or soil.


Evolution

Owlflies appear to have evolved from a common ancestor with Stilbopterygidae. These, in turn, evolved from a common ancestor with Palparidae, which evolved from a common ancestor with the true antlions, or
Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the larvae ...
.


Taxonomy and etymology

The family Ascalaphidae was first described by the French entomologist
Jules Pierre Rambur Jules Pierre Rambur (21 July 1801 – 10 August 1870) was a French entomologist. Rambur was born in Chinon. He studied the insect fauna of Corsica and Andalusia. He was the author of ''Histoire naturelle des insectes'' (1842) amongst other works ...
in 1842. The name is from Greek ''askalaphos'', a kind of owl. In
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the Cosmogony, origin and Cosmology#Metaphysical co ...
, Ascalaphos was the custodian of the orchard of
Hades Hades (; grc-gre, ᾍδης, Háidēs; ), in the ancient Greek religion and myth, is the god of the dead and the king of the underworld, with which his name became synonymous. Hades was the eldest son of Cronus and Rhea, although this also ...
, god of the underworld; the goddess
Demeter In ancient Greek religion and mythology, Demeter (; Attic: ''Dēmḗtēr'' ; Doric: ''Dāmā́tēr'') is the Olympian goddess of the harvest and agriculture, presiding over crops, grains, food, and the fertility of the earth. Although s ...
transformed him into an owl.


Fossil history

The owlflies are known from
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 ...
s of adults and larvae, often encased in
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
. Most of these cannot be placed in a particular subfamily. Most are known from the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
. The
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
'' Mesascalaphus'' was thought to be a more basal member of the family, but it is now believed to be a member of Mesochrysopidae.


Phylogeny

Total evidence analysis (several genes + morphology) in 2019 recovered Ascalaphidae as
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
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. This followed nuclear phylogenomic analysis in 2018, which recovered Ascalaphidae as a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
lineage within Myrmeleontidae; authors in that paper therefore sank Ascalaphidae into the Myrmeleontidae as subfamily Ascalaphinae, put the Stilbopterygini and the Palparini into the Ascalaphinae, and reduced its subfamilies to the rank of tribes. Molecular analysis in 2018 using
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
l
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosoma ...
and
mitogenomic Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
data also placed the Ascalaphidae as sister to the
Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the larvae ...
as the most advanced groups within the Neuroptera. The
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 ...
record has contributed to an understanding of the group's phylogeny.Parker, S. P. (ed.), 1982: Synopsis and classification of living organisms. Vols. 1 & 2. McGrew-Hill Book Company The phylogeny of the owlflies has remained uncertain, with many of the higher taxa apparently not natural groups (
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s).


External

Neuropteran subfamilies are described in Winterton and colleagues 2017 and Jones 2019.


Internal

Machado et al 2018 proposes a classification below
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
level, into
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...
(names ending with –ini): Groups formerly considered part of "Myrmeleontidae" are underscored and marked "Myrm." Jones 2019 presents a total-evidence phylogeny, preferring to classify only to family level:


References


External links

*
Owlflies on Stamps
{{Authority control Neuroptera Neuroptera families Extant Oligocene first appearances