Ascalaphidae In Parappa
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Ascalaphidae is a family of
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s in the order
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
, 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 (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
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 their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...
s; some of them make use of
self-decoration camouflage Self-decoration camouflage is a method of camouflage in which animals or soldiers select materials, sometimes living, from the environment and attach these to themselves for concealment. The method was described in 1889 by William Bateson, who o ...
.


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 dragonflies are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
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 (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. ...
) have short, weakly clubbed antennae, smaller eyes, and reticulate
wing venation Insect wings are adult outgrowths of the insect exoskeleton that enable insect flight, insects to fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often ...
. 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 found ...
such as ''
Ululodes ''Ululodes'' is a genus of owlflies in the tribe Ululodini. There are at least 27 described species in ''Ululodes''. Etymology For comments on etymology, see Jones & Oswald (2013) Species As of 2024 the genus contains: * '' Ululodes apolli ...
'' have large divided eyes and
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
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 in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
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. Description The adult is a large insect somewhat resembling a dragonfly. Its body, eyes, and l ...
'', are active during the day.


Anti-predator defences

When disturbed, some owlflies release a strong,
musk Musk 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 substances with similar odors. ' ...
-like chemical to deter enemies. The
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
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 anima ...
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 (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e are
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...
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 (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. ...
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 Self-decoration camouflage is a method of camouflage in which animals or soldiers select materials, sometimes living, from the environment and attach these to themselves for concealment. The method was described in 1889 by William Bateson, who o ...
.
Pupation A pupa (; : 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 their life cycle, the stages th ...
occurs in a spheroidal
silk cocoon Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is most commonly produced by certain insect larvae to form cocoons. The best-known silk is obtaine ...
in
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
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 (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. ...
.


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 Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
, Ascalaphos was the custodian of the orchard of
Hades Hades (; , , later ), in the ancient Greek religion and Greek mythology, mythology, is the god of the dead and the king of the Greek underworld, underworld, with which his name became synonymous. Hades was the eldest son of Cronus and Rhea ...
, god of the underworld; the goddess
Demeter In ancient Greek religion and Greek mythology, mythology, Demeter (; Attic Greek, Attic: ''Dēmḗtēr'' ; Doric Greek, Doric: ''Dāmā́tēr'') is the Twelve Olympians, Olympian goddess of the harvest and agriculture, presiding over cro ...
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 preserve ...
s of adults and larvae, often encased in
Baltic amber Baltic amber or succinite is amber from the Baltic region, home of its largest known deposits. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that this forested region provided the re ...
. Most of these cannot be placed in a particular subfamily. Most are known from the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
. The
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
'' Mesascalaphus'' was thought to be a more basal member of the family, but it is now believed to be a member of
Mesochrysopidae Mesochrysopidae is an extinct family of lacewings known from the Jurassic and Cretaceous periods. They are considered to be closely related to green lacewings of the family Chrysopidae. The family are also alternatively considered a Paraphyly, p ...
.


Phylogeny

Total evidence analysis (several genes + morphology) in 2019 recovered Ascalaphidae as
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
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 refuted nuclear phylogenomic analysis in 2018, which recovered Ascalaphidae as a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
lineage within Myrmeleontidae. Molecular analysis in 2018 using
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
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 ribosomal ...
and mitogenomic 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 (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. ...
as the most advanced groups within the
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
. 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 preserve ...
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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
level, into
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
(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