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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 The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in the ...
, 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 dayli ...
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 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 true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threaten ...
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 l ...
) have short, weakly clubbed antennae, smaller eyes, and reticulate wing venation. 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. References Myrmeleontidae Inse ...
such as ''
Ululodes ''Ululodes'' is a genus of owlflies in the tribe Ululodini. There are 26 described species in ''Ululodes''. Species * ''Ululodes apollinaris'' * ''Ululodes arizonensis'' * ''Ululodes banksi'' * ''Ululodes bicolor'' * '' Ululodes brachyceru ...
'' 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 dayli ...
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 ...
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 ...
'', 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 s ...
-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 tors ...
in ''Ululodes quadrimaculatus'' is raised at rest,
mimicking In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry ...
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) o ...
, exposing a strongly-contrasting patch of pale colour (white or cream), either as a deimatic display 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. Th ...
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 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 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 larva ...
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 ( 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 the ...
occurs in a spheroidal silk cocoon 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 l ...
.


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 Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of ...
, 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 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
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 ...
'' Mesascalaphus'' was thought to be a more
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
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 paraphyletic ...
.


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 ...
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 mitochondrial
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 riboso ...
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 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 l ...
as the most advanced groups within the
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in the ...
. 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 ( clades).


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 group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
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 confl ...
(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