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

and see text

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

[edit]

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

[edit]
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

[edit]
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

[edit]
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

[edit]

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

[edit]

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

[edit]

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

[edit]

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

[edit]

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

[edit]

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

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

Myrmeleontoidea

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Owlfly

View on Grokipedia
from Grokipedia
Owlfly is the for belonging to the Ascalaphinae in the family Myrmeleontidae (antlions) within the order , comprising approximately 450 extant species distributed worldwide, primarily in tropical and subtropical regions. Formerly classified as a separate family Ascalaphidae, recent phylogenetic studies as of 2019 have reclassified them within Myrmeleontidae. These fast-flying, crepuscular or nocturnal predators superficially resemble dragonflies due to their slender bodies, large compound eyes, and transparent wings with spans of 40–80 mm, but are distinguished by their long, clubbed antennae and raptorial mouthparts adapted for capturing prey in flight. Adult owlflies are agile aerial hunters that primarily feed on small flying such as flies and midges, often active at and dawn, while resting during the day with their abdomens raised on . Their larvae, in contrast, are predators resembling those of antlions, with flattened, camouflaged bodies up to 20 mm long equipped with powerful, toothed mandibles for seizing mites, eggs, larvae, and small arthropods in leaf litter or on tree bark. The life cycle is holometabolous, featuring egg clusters laid on twigs—often guarded by a silken shield or trophic eggs to provision hatchlings—followed by three larval instars that pupate in silken cocoons within or debris, with adults emerging in warmer months. Owlflies play a significant ecological role as biological control agents, devouring vast numbers of pest , and are found across diverse habitats from arid Australian outback to North American woodlands, though they are more abundant in the . Some species exhibit defensive behaviors, such as releasing a noxious when handled, and the includes approximately 100 genera.

Description

Morphology

Adult owlflies possess a slender, elongated body typically measuring 25–50 mm in length, with large, bulging eyes that occupy much of the head and confer an owl-like appearance. These eyes are a prominent feature, often divided by a transverse sulcus in genera such as Ululodes, while remaining undivided in genera like Ascaloptynx. The antennae are long and clubbed, inserted laterally on the head and nearly as long as the body, with the club often bi-colored at the tip. The wings are broad and transparent, with spans of 40–80 mm, featuring prominent venation and a lacy appearance; they are typically held roof-like over the body at rest, with hindwings sometimes shorter than the forewings. Larvae exhibit a stout, flattened, oval body that is in shades of gray, brown, or black, often with a fringe of tiny fin-like lateral extensions on the and . They possess powerful, caliper-like mandibles equipped with multiple teeth for prey capture and can actively decorate their dorsum with sand and debris for additional . Sexual dimorphism is evident in many species, with males generally featuring larger eyes and more pronounced antennal clubs, including specialized setae on the antennae that are less developed in females.

Distribution and Habitat

Owlflies (family Ascalaphidae) are distributed worldwide but predominantly in tropical and subtropical regions, with approximately 450 species recorded globally. They are largely absent from cold temperate zones but occur marginally in warmer parts of some temperate areas, reflecting their preference for warmer climates. Highest occurs in the Afrotropical, Oriental (including Indo-Malayan), and Australasian regions, with around 150 species in alone and about 40 species in . In contrast, hosts only eight species north of , primarily in the genus Ululodes, which are restricted to the . These favor warm, dry to semi-arid environments such as savannas, open woodlands, and scrublands, where adults typically perch on during their crepuscular or nocturnal activity periods. Larvae inhabit leaf , soil, or under bark, ambushing prey in these ground-level microhabitats. Such preferences align with predation hotspots in open areas, facilitating their hunting strategies. Regional is pronounced, with high turnover in the Indo-Malayan and Afrotropical realms contributing to overall diversity. For instance, harbors at least seven , several of which are island endemics confined to forested areas. Altitudinally, owlflies range from to montane forests up to 2,000 meters, as observed in Neotropical populations.

Biology

Life Cycle

The life cycle of owlflies (family Ascalaphidae) consists of four distinct stages: egg, larva, pupa, and adult, typical of holometabolous in the order . Females lay eggs in clusters on such as grass stems or twigs, often numbering 50 to 100 per mass; these eggs may be positioned on stalks called rapagula in some or surrounded by protective structures like trophic eggs or barriers to deter predation by and other ground-dwelling arthropods. In some , first-instar larvae form a defensive ring around the egg mass before dispersing. The lasts approximately 3 to 4 weeks, varying with temperature and , after which first-instar larvae emerge. Larval development spans three instars and is characterized by campodeiform morphology—elongated, active bodies with well-developed legs—enabling an predation on small arthropods. The larval stage typically endures 6 to 12 months, influenced by and food availability; in temperate regions, larvae overwinter in within soil or leaf litter, resuming growth in spring. Larval , often involving debris coverage, enhances survival during this prolonged phase. Upon reaching maturity, third-instar larvae construct silk cocoons in , leaf litter, or under bark for pupation; this non-feeding stage lasts 2 to 4 weeks, during which the undergoes . Eclosion occurs when the splits the cocoon longitudinally, allowing the to emerge. Adults are short-lived, surviving a few weeks, with their primary focus on rather than feeding, though they may consume or prey opportunistically. involves chemical signaling via pheromones and visual displays, such as aerial pursuits or perch presentations, often at or night in crepuscular species. varies geographically: most temperate populations are univoltine, completing one generation per year, while tropical may be multivoltine, producing two or more broods annually within a 12-month cycle.

Predation and Diet

Adult owlflies are primarily crepuscular or nocturnal aerial predators, with some species active during the day, that capture small flying , such as flies, moths, and midges, in mid-flight using their acute vision and agile maneuvers. They perch on during the day and actively hunt at or dawn, employing swift, direct flights to intercept prey. Feeding occurs primarily in warmer seasons when activity levels peak, aligning with increased availability. Owlfly larvae function as ambush predators in leaf litter, on tree bark, or , targeting ground-dwelling arthropods including , beetles, and other small . They employ a sit-and-wait strategy, camouflaged by debris and bristles, and seize prey with oversized mandibles that open up to 180 degrees. Upon capture, larvae inject through their mandibles to liquefy prey tissues via extraoral digestion, then suck up the resulting fluids. This process is voracious and continuous except during molts, with larvae opportunistically feeding year-round in suitable microhabitats. In their ecosystems, owlflies occupy a trophic position as mid-level to apex predators within microhabitats, regulating populations of pest like fruit flies and contributing to natural . Their predatory habits, supported by specialized mouthparts and eyes, underscore their role in maintaining ecological balance.

Anti-predator Adaptations

Owlfly larvae employ sophisticated strategies to evade detection by predators, primarily through self-decoration with environmental . Many possess specialized bristle-like structures called dolichasters that adhere , particles, , or matter to their dorsum, enabling them to mimic twigs, rocks, or surrounding substrate. For instance, in European such as Ascalaphus festivus and Libelloides longicornis, these dolichasters retain fine rock dust or , allowing larvae to blend seamlessly into sandy or rocky habitats where they remain motionless as ambush predators. This -carrying adaptation not only conceals the larvae but also enhances their during extended periods of inactivity on the surface or under leaf litter. owlflies similarly rely on cryptic coloration, with their slender bodies and wings often patterned in shades of brown, gray, or to match vegetation; some further mimic broken twigs by adopting a rigid, elongated posture while at rest. To actively deter approaching threats, owlflies utilize deimatic displays that startle predators through sudden visual contrasts. In species like Haploglenius luteus, adults (particularly males) rapidly flex a hinged flap on the pronotum to expose a brilliant patch, creating a flash of contrasting color against their otherwise drab . This brief revelation aims to interrupt a predator's attack, providing a momentary escape opportunity, and may be reinforced by underlying distastefulness, as some ascalaphids produce unpalatable secretions. Complementing these visual tactics, certain larvae possess chemical defenses. Behavioral adaptations further bolster owlfly survival, particularly in adults, which exhibit powerful, swift flight capabilities—the strongest among —to execute rapid evasions. Many species are crepuscular or nocturnal, minimizing encounters with diurnal predators like birds while at or dawn. When disturbed, adults often release a strong, musk-like or noxious odor from glandular secretions, deterring handling by vertebrates. These combined strategies contribute to effective predator avoidance, with larval and stillness supporting high success, while adult mobility and displays enable evasion in open habitats.

Taxonomy and Evolution

Etymology and Classification History

The common name "owlfly" derives from the insects' large, prominent eyes that resemble those of an , evoking a nocturnal predatory gaze despite their diurnal habits. The family name Ascalaphidae originates from the type genus Ascalaphus, which in turn stems from the Greek word askalaphos, referring to a type of owl and drawing from where Ascalaphus was a figure associated with the . Early taxonomic work on owlflies began in the early , with species initially classified alongside antlions in the broader group of Myrmeleonidae due to superficial similarities in larval morphology and predatory behavior. The family Ascalaphidae was formally established in 1842 by French entomologist Jules Pierre Rambur in his Histoire naturelle des insectes névroptères, distinguishing owlflies based on their unique combination of features and antennae. During the mid- to late , British entomologist Robert McLachlan contributed significantly by describing and expanding several genera, such as Bubopsis in 1898 and Glyptobasis in 1873, which helped delineate owlfly diversity in the tropics. The subfamily Haplogleniinae was defined in 1853 by Edward Newman, characterized by entire eyes lacking a transverse suture, marking an early subdivision within the family. In the , key advancements included Herman van der Weele's comprehensive 1909 Ascalaphiden monographisch bearbeitet, which systematically reviewed and illustrated numerous , primarily from and , laying the groundwork for subsequent revisions. This work highlighted morphological variations and spurred debates over the familial status of Ascalaphidae, with some classifications treating it as a distinct family sister to Myrmeleontidae, while others proposed subfamily integration based on shared larval traits. Recent revisions, building on these foundations, recognize approximately 450 across nearly 100 genera, reflecting ongoing refinements in amid phylogenetic reevaluations.

Current Taxonomy

Owlflies are traditionally classified in the Ascalaphidae within the superfamily Myrmeleontoidea of the order , encompassing approximately 450 species distributed across nearly 100 genera worldwide. As of 2024, approximately 435-450 species are recognized. However, a significant taxonomic has emerged since 2019, when phylogenomic analyses proposed synonymizing Ascalaphidae with the antlion family Myrmeleontidae, thereby demoting owlflies to the Ascalaphinae; this revision is supported by anchored hybrid enrichment data from 325 genes, highlighting owlflies as derived s, though traditional family status persists in some regional checklists. The family is currently divided into five recognized subfamilies, primarily distinguished by morphological traits such as eye structure and wing venation, with distributions often reflecting biogeographic patterns: Ascalaphinae (cosmopolitan, including many species), Ululodinae (predominantly Neotropical, featuring split-eyed forms), Albardiinae (Afrotropical and Oriental), Haplogleniinae (mainly Afrotropical), and Melambrotinae (Oriental and Australasian). These subfamilies accommodate the family's diversity, with Ascalaphinae being the most speciose and widespread. Prominent genera include Ululodes, which dominates in with species adapted to arid habitats, Ascalaphus in the Palaearctic region known for its nocturnal habits, and Deleproctophylla in , characterized by elongated antennae and high . Recent taxonomic work has added to this roster, such as the 2024 revision of the Indian genus Glyptobasis, which described new species and clarified synonymies in the Oriental . Taxonomic challenges persist due to the group's high , particularly in , resulting in numerous undescribed species and incomplete inventories; for instance, barcode-assisted checklists from the in 2024 revealed new distribution records and genetic insights for regional diversity. Revisions remain fragmentary in the tropics, where loss exacerbates documentation gaps, though species counts have held steady at around 450, bolstered by 10-20 new descriptions since 2020, including novel taxa from and .

Fossil Record

The fossil record of owlflies (family Ascalaphidae) is sparse compared to other neuropteran families, primarily due to the soft-bodied nature of their larvae and the aerial habits of adults, with most preservations occurring in amber deposits that favor tropical and subtropical environments. The earliest known fossils are larval specimens from mid-Cretaceous (Albian-Cenomanian) Burmese amber, dating to approximately 100 million years ago, including several specimens exhibiting debris-carrying camouflage behaviors similar to extant forms. These rare fragments highlight early diversification within the family, though adult fossils from this period remain unknown. Eocene deposits provide the first evidence of adult owlflies, with wing imprints and partial specimens showing primitive venation patterns. A notable example is Pseudoameropterus ambiguus, a new genus and species described from compression fossils in the middle Eocene (approximately 46 million years ago) Kishenehn Formation of northwestern , , representing one of the oldest definitive adult records and indicating North American presence during a period of global warming. Eocene (about 44 million years ago) yields more complete specimens, including larvae and adults with features akin to modern Ascaloptynx-like forms, though specific species counts from early descriptions total around seven for the broader myrmeleontiform group. Oligocene and Miocene ambers further document owlfly diversity, with preservation biases toward larvae in resin. Oligocene Baltic amber contains additional adult and larval inclusions, while Miocene Dominican amber preserves at least eight larval specimens, characterized by large, three-toothed stylets and ambush-predatory morphology. A recently described larva from Miocene (approximately 16–23 million years ago) Mexican amber in further expands the New World record, featuring prominent eye hills and setose trunk protrusions. Overall, around 38 fossil owlfly larval specimens have been documented across these deposits, spanning roughly 20 described , with extinct lineages like early haplogleniine forms suggesting origins in tropical paleoforests and gradual size reduction in some clades post-Eocene. These fossils imply resilience to climatic shifts without evident mass extinction effects, contributing to understandings of neuropteran .

Phylogeny

Owlflies (Ascalaphidae) occupy a derived position within the superfamily Myrmeleontoidea of the order , emerging as antlion-like ancestors and nesting closely with the family Myrmeleontidae (s). A phylogenomic using anchored hybrid enrichment of 325 genes across 207 demonstrated that owlflies are embedded within Myrmeleontidae, rendering the traditional separation of Ascalaphidae paraphyletic and supporting their synonymy under an expanded Myrmeleontidae. This placement aligns owlflies as sister to core antlion lineages, with Nemopteridae (thread-winged lacewings) as the immediate outgroup to the combined Myrmeleontidae + former Ascalaphidae , reinforcing the of Myrmeleontoidea. Internally, the relationships among owlfly subfamilies exhibit strong for most groups in total-evidence analyses, though some conflicts persist. A 2019 study integrating 25 morphological characters with molecular data from 76 exemplars (including 57 owlfly species) via parsimony, Bayesian, and maximum-likelihood methods positioned Haplogleniinae as the basal subfamily within Ascalaphidae, with and African/Malagasy lineages forming well-supported monophyletic groups. Ululodinae emerged as monophyletic across analyses but showed placement inconsistencies: sister to Myrmeleontidae in DNA-only maximum-likelihood trees, yet integrated within Ascalaphidae in total-evidence frameworks. Traditional subfamilies like Ascalaphinae and Haplogleniinae were not monophyletic in all analyses, prompting a revised higher-level classification into tribes such as Ululodini and Haplogleniini. Molecular evidence further bolsters these ties, with mitochondrial genome sequencing confirming close affinities to Myrmeleontidae. The complete mitogenome of Suhpalacsa longialata (15,911 bp), analyzed via and maximum likelihood, clustered as sister to other owlflies and positioned non-monophyletic Ascalaphidae within Myrmeleontidae, highlighting gene rearrangements like tRNA transposition as shared traits. DNA barcode data, particularly COI sequences, have aided delimitation by generating reference libraries; for instance, a 2024 barcode-assisted checklist for Georgian owlflies produced five high-quality barcodes from 128 specimens across four , facilitating identification and first records like Deleproctophylla meridionalis. In the broader phylogeny, Myrmeleontoidea (including owlflies) diverges early from lineages like Dilaridae (pleasing lacewings) and Hemerobiidae (brown lacewings), with the former basal to Cysneuroptera and the latter in Hemerobioidea. Divergence time estimates place the split of Myrmeleontidae and Ascalaphidae ancestors in the mid-Late to , around 100–150 million years ago, coinciding with the emergence of modern diversity. The debate over owlfly status as a distinct versus a subfamily (Ascalaphinae) remains unresolved, with 2024 morphological studies on male genitalia reinforcing phylogenomic evidence for integration into Myrmeleontidae through shared synapomorphies like the gonarcus structure, though practical retention of Ascalaphidae persists in some taxonomic works for clarity.

References

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