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Mantispidae

Mantispidae, commonly known as mantidflies or mantispids, is a family of predatory insects belonging to the order Neuroptera, distinguished by their superficial resemblance to praying mantises through an elongated prothorax and raptorial forelegs adapted for grasping prey.^[1] Adults typically feature clear, veined wings that fold flat over the abdomen at rest, a mantis-like head with large compound eyes, and chewing mouthparts, with body lengths ranging from about 5 to 47 mm and wingspans of 5 to 30 mm.^[2] The family encompasses approximately 370 extant species across 42 genera, divided into three subfamilies: Drepanicinae, Calomantispinae, and Mantispinae.^[3] These insects exhibit a worldwide distribution, excluding Antarctica, with the highest species diversity in the Neotropics, Australia, and the Afrotropics, reflecting a Gondwanan biogeographic origin.^[4] Larvae undergo hypermetamorphosis, a specialized form of complete metamorphosis where the first instar is highly mobile and planidial, actively seeking out hosts such as spider egg sacs or the nests of solitary bees and wasps, while later instars become more sedentary and endoparasitic or kleptoparasitic within those hosts.^[1] Adults are generalist predators, primarily active at dusk or nocturnally, feeding on small arthropods like aphids or other soft-bodied insects using their raptorial forelegs, and they may also consume nectar or sap; some species, such as Climaciella brunnea, exhibit wasp mimicry for protection.^[4]^[1] Despite their predatory habits, mantidflies pose no threat to humans and play a beneficial role in natural pest control.^[5]

Description

Adult Morphology

Adult mantispids are small to moderate-sized insects, with body lengths typically ranging from 10 to 30 mm.^[6] Their coloration is often cryptic, featuring shades of brown, gray, or yellow, which aids in camouflage; some species exhibit patterns mimicking wasps or ants for protective resemblance.^[7]^[6] The head is triangular in shape, featuring large compound eyes that provide wide visual fields, and prominent ocelli in certain species.^[7]^[8] Antennae vary from filiform to pectinate or moniliform types, with sexual dimorphism evident in some genera where males possess larger antennal clubs or more elaborate pectinate structures. Mouthparts are of the chewing type, with strong mandibles and maxillae adapted for both predation on small insects and nectar or pollen feeding.^[9] The thorax is characterized by an elongated prothorax, which supports the raptorial forelegs and contributes to the mantis-like appearance.^[10] These forelegs are enlarged and spiny, functioning as grasping appendages; the coxa is robust and attached to the prothorax, followed by a short trochanter, a thickened femur bearing prominent spines, and a tibia with opposing spines that folds against the femur to secure prey.^[8]^[9] The middle and hind legs are ambulatory, similar to those of other neuropterans. Wings consist of two pairs of similar, membranous structures with a net-like venation pattern typical of Neuroptera, including a well-defined pterostigma and multiple radial sectors; at rest, they are typically held roof-like over the abdomen.^[9] Wing spans range from 10 to 30 mm, varying by species.^[11] The abdomen is slender and elongated, comprising up to 10 segments, with females in some subfamilies possessing a short ovipositor for egg deposition.^[9] Overall, these features distinguish mantispids from other neuropterans through their mantis-mimicking raptorial forelegs and elongated prothorax.^[10]

Immature Morphology

The immature stages of Mantispidae exhibit hypermetamorphosis, with larvae passing through three distinct instars adapted for a predatory lifestyle. The first instar, often termed the planidium, is campodeiform—dorso-ventrally flattened and prognathous—with a sclerotized exoskeleton enabling rapid, agile movement; body length typically ranges from 0.5 to 1 mm at hatching.^[12] Later instars shift to a more grub-like or scarabeiform morphology, becoming sedentary with reduced sclerotization and body lengths reaching 7-10 mm in the third instar.^[13]^[1] Sensory structures in mantispid larvae support navigation and host location. The first instar features short, three-segmented antennae and a pair of lateral stemmata—simple, single-lens eyes positioned on each side of the head—for basic visual detection.^[14] Tactile setae are distributed across the body, particularly on the head and thorax, aiding in sensory perception during movement over substrates like silk or soil.^[15] The mouthparts include prominent, curved, hollow mandibles and maxillae forming needle-like stylets for piercing and fluid extraction, such as hemolymph or liquefied egg contents, rather than purely chewing.^[12] Legs in immature mantispids are ambulatory rather than raptorial, reflecting their non-jumping locomotion. In the first instar, legs are long and well-developed, comprising coxa, trochanter, femur, and a tibia-tarsus fusion ending in paired claws and a trumpet-shaped empodium or pseudoclaw for secure attachment to hosts or surfaces.^[12]^[16] Subsequent instars have shorter, less prominent legs suited to their stationary feeding position within host structures.^[12] The pupal stage is exarate, with legs, wings, and antennae free from the body, allowing limited mobility as the pharate adult can "walk" the pupa to an emergence site.^[17] Pupation occurs within a silken cocoon, often spun inside the remnants of the third larval skin or host egg sac, lasting about one month.^[13] This stage transitions the immobile larva to the winged adult, with the pupa typically measuring approximately 7-10 mm in length.^[13]

Distribution and Ecology

Global Distribution

Mantispidae exhibit a nearly cosmopolitan distribution, occurring on all continents except Antarctica and showing a clear preference for tropical and subtropical climates, where the majority of species diversity is concentrated. The family is absent from polar regions due to their intolerance of extreme cold, but representatives are found across a wide latitudinal range from equatorial zones to temperate areas. This broad yet uneven global presence reflects the group's adaptation to warmer environments, with approximately 400 species documented worldwide.^[4]^[18] Species richness varies significantly by biogeographic region (as of 2012), underscoring patterns of tropical diversification. The Oriental region hosts the highest number at approximately 121 species, followed closely by the Afrotropical (99 species) and Neotropical (95 species) regions, which together account for a substantial portion of global diversity. In contrast, the Australasian region has around 67 species, while the temperate Palearctic (33 species) and Nearctic (19 species) regions support fewer taxa, highlighting a gradient of decreasing abundance toward higher latitudes. These figures, based on taxonomic inventories from 2012, indicate ongoing discoveries, particularly in understudied tropical areas; recent studies continue to describe new species, especially in the Neotropics.^[18]^[19] Endemism is pronounced in key tropical biodiversity hotspots, contributing to regional uniqueness. In the Neotropics, the Amazon Basin stands out as a center of high endemism, with numerous species restricted to its rainforests. Similarly, Madagascar harbors endemic genera such as Madantispa (three species), illustrating island isolation's role in speciation. While island endemics are notable, these patterns emphasize the family's reliance on stable, biodiverse habitats.^[18]^[20] The historical biogeography of Mantispidae points to an origin in Gondwana during the Jurassic period, around 150–200 million years ago, serving as the epicenter for early radiation. Fossil evidence supports this, with Jurassic specimens like Liassochrysa stigmatica from the Lower Jurassic of Dobbertin, Germany, indicating an ancient lineage. Subsequent diversification led to spread into Laurasia, as evidenced by Mesozoic amber inclusions from Cretaceous deposits in Myanmar and Spain, revealing the family's presence in northern supercontinental fragments by the mid-Cretaceous. Introductions remain rare, though a few species have become adventive in non-native areas through human activities, such as trade and transport in controlled environments.^[20]^[21]^[22]

Habitat and Ecological Interactions

Mantispidae species primarily inhabit diverse terrestrial environments such as forests, grasslands, and scrublands, where vegetation provides suitable perches for adults and access to prey.^[23] Adults are commonly observed in the lower to middle layers of vegetation, ambushing small arthropods during daylight or nocturnal hours.^[6] Larval stages, in contrast, are found in concealed microhabitats including soil, leaf litter, and especially spider egg sacs or retreats, reflecting their dependence on spider hosts for development.^[24] Microhabitat selection is closely tied to the availability of spider populations, as first-instar larvae actively seek out and board female spiders to access egg sacs, often in proximity to web-building or hunting spider colonies.^[4] Certain species, such as those in the subfamily Mantispinae, occur in arid or semi-arid regions like the Texas Hill Country, where their activity peaks seasonally during periods of higher prey abundance and milder conditions.^[1] In ecosystems, mantispids serve as key predators that regulate populations of small insects and spiders; larvae consume spider eggs and immatures, potentially curbing outbreaks of web-building arachnids, while adults prey on a range of arthropods including flies and caterpillars.^[24] Adults also visit flowers to hunt pollinating insects, incidentally aiding pollination through nectar visitation, though their primary diet remains predatory.^[25] Mantispids engage in defensive interactions via Batesian mimicry, with many adults resembling hymenopterans such as wasps in coloration and body form to avoid predation by birds and other vertebrates.^[4] They may face indirect competition with other neuropteran families, like antlions, for shared prey resources in vegetated habitats, though such overlaps are not well-documented.^[6] Conservation threats to mantispids are generally minor, with habitat fragmentation and loss in tropical forests impacting species diversity by reducing spider host availability; however, no species are classified as endangered as of 2025.^[26] Seasonal life history varies geographically: multivoltine cycles prevail in tropical areas, allowing multiple generations annually, whereas temperate populations are largely univoltine, with adults emerging from late spring through autumn.^[6]

Life Cycle

Reproduction and Development

Mating in Mantispidae typically involves courtship displays where males use their raptorial forelegs to grasp females, accompanied by wing fanning and abdominal vibrations to attract mates.^[27] In species such as Mantispa uhleri, males adopt a "wings-up" stance and engage in reciprocal foreleg sparring before copulation, which lasts approximately 23–36 minutes.^[27] A sweetish odor, possibly a male pheromone, is often detected during these interactions, while in Mantispa viridis, males evert intertergal membranes that may release recognition pheromones.^[28]^[27] Females oviposit clusters of 100–3,000 eggs, often attached to vegetation or substrates via short silken stalks, though some species deposit them near or directly on spider webs to facilitate larval access to hosts.^[29]^[27] Egg morphology is generally elongate and ellipsoid, measuring about 0.36 mm in length and 0.18 mm in width, with a tannish-yellow to light pink coloration and ribbed or banded surface in some cases.^[27] Mated females produce multiple clutches over several days; for example, M. uhleri averages ~13 clutches of 600–3,000 eggs each (all fertile), while M. viridis produces up to ~59 clutches with ~23 fertile (~800 eggs each); unmated females lay infertile eggs.^[28]^[27] Eggs incubate for 1–2 weeks under favorable conditions, hatching into highly mobile, campodeiform first-instar larvae that are approximately 0.88 mm long and actively seek hosts.^[27]^[28] Hatching success varies by species (e.g., 84% in M. viridis, 98% in M. uhleri), with eclosion occurring synchronously within clutches after visible embryonic development, such as eye formation around day 5–7.^[28]^[27] Mantispidae undergo complete metamorphosis with hypermetamorphosis and three larval instars, transitioning from active first instars to more sedentary later stages before pupation in a cocoon.^[12] The total developmental timeline from egg to adult varies by species and climate, spanning ~1 month in laboratory conditions for M. uhleri (28 days lab, 37 days field) or up to 1 year in temperate regions for species like Climaciella brunnea due to diapause or overwintering as larvae.^[29]^[27] Parental care is absent in Mantispidae, with eggs and larvae left unattended after oviposition. Some species exhibit delayed hatching or diapause to synchronize larval emergence with spider host availability, enhancing survival rates. Voltinism varies from 1–3 generations per year, influenced by latitude and climate, with tropical species potentially producing more generations than those in temperate zones.^[29]

Larval Predation Strategies

Mantispid larvae exhibit diverse predation strategies, primarily as active predators or parasitoids targeting spider eggs, guided by chemosensory cues such as spider silk and excreta deposited on substrates. First-instar larvae, which are campodeiform and highly mobile, detect these chemical signals to locate potential hosts, raising their bodies in a phoretic posture to board passing spiders or burrow directly into exposed egg sacs. This behavior allows them to exploit spider reproductive efforts, with larvae in the subfamily Mantispinae specializing in internal feeding on eggs and embryos once inside the sac.^[30] In spider egg predation, prevalent among Mantispinae, a single first-instar larva penetrates the egg sac—either by boarding the female spider prior to oviposition or directly invading completed sacs—and consumes up to 100 eggs or developing embryos over its development. The larva pierces eggs with modified mandibles and maxillae forming a sucking tube, draining contents while often halting further spider embryo development through chemical inhibition rather than mechanical damage. This internal parasitoidism ensures a nutrient-rich environment, with the larva growing substantially through subsequent instars within the sac. Representative examples include species like Mantispa uhleri, which target hunting spiders such as wolf spiders, consuming entire clutches in sacs containing dozens to hundreds of eggs.^[31] Ectoparasitoidism occurs when larvae attach to female spiders, feeding on hemolymph by piercing the exoskeleton while awaiting egg sac production, sometimes sustaining themselves for months in book lungs or near the pedicel. Larvae can transfer hosts opportunistically, such as during spider mating or cannibalism events, allowing access to new egg batches and enhancing survival odds. In basal subfamilies like Symphrasinae, predation shifts to alternative hosts, with larvae acting as ectoparasitoids or kleptoparasites on nest-building Hymenoptera, targeting larvae and pupae in wasp or bee nests rather than spiders.^[4] Instar progression reflects these strategies: early instars (first) are active hunters, mobile and phoretic to secure hosts, while later instars (second and third) become scarabaeiform, less mobile, and remain embedded in the egg sac or host, feeding until pupation occurs within the sac or adjacent silk. Pupation typically happens nearby if the host is depleted, with the pupa enclosed in a silken cocoon. Survival rates are low, with high larval mortality due to spider host defenses such as grooming, palp palpation, or outright consumption of attached larvae; adaptive behaviors, including positioning in less accessible body regions, mitigate some risks but do not eliminate the challenges posed by vigilant hosts.^[12]

Taxonomy and Systematics

Classification History

The family Mantispidae was formally established by William Elford Leach in 1815 as part of the order Neuroptera, recognizing the distinctive raptorial forelegs and mantis-like appearance of its members, with early species descriptions noting similarities to praying mantises dating back to the mid-18th century.^[32]^[24] Initial taxonomic placements varied, often grouping them with Orthoptera due to superficial resemblances, before their firm assignment to Neuroptera.^[33] During the 19th and early 20th centuries, the classification evolved with the erection of subfamilies by Anton Handlirsch in 1906, who outlined early divisions based on wing venation and leg morphology, followed by revisions from Robin John Tillyard in 1925 that refined generic groupings within the family.^[34]^[35] A major advancement came in 1986 when Kevin J. Lambkin conducted a comprehensive revision of the Australian fauna, proposing a classification into four subfamilies—Symphrasinae, Drepanicinae, Mantispinae, and Calomantispinae—based on phylogenetic analysis of morphological characters such as prothoracic structure and genital features.^[36] Subsequent key revisions included Michael Ohl's 2002 monograph on Symphrasinae, which provided detailed morphological and phylogenetic insights into this subfamily's relationships and diversity, emphasizing character evolution in raptorial appendages. Recent molecular studies, such as Wang et al. (2017), have utilized mitochondrial genomes to confirm the monophyly of Mantispidae within Neuroptera, supporting its placement in the Hemerobiiformia clade while highlighting close affinities to other mantispaoid families like Berothidae and Rhachiberothidae.^[37] Recent studies (e.g., Ardila-Camacho et al., 2021; 2024) have proposed reclassifying Symphrasinae within Rhachiberothidae, challenging the traditional four-subfamily structure of Mantispidae, though some sources retain the inclusion as of 2025.^[3] Challenges persist, including nomenclatural ambiguities in the Neotropical fauna, which were addressed in a 2025 study resolving synonymies and type designations for numerous species through examination of historical collections.^[19] Currently, Mantispidae comprises approximately 400 extant species across more than 40 genera, with ongoing debates regarding the phylogenetic inclusion of certain subfamilies like Symphrasinae, potentially aligning them closer to Rhachiberothidae based on combined morphological and molecular evidence.^[38] Fossils have been integrated into phylogenies to trace evolutionary origins, reinforcing the family's position as sister to other raptorial lineages in Mantispoidea, though non-monophyly hypotheses continue to be tested.^[9]

Extant Subfamilies

The family Mantispidae comprises approximately 400 extant species distributed among four subfamilies, with diversity concentrated in tropical regions worldwide.^[38] These subfamilies are distinguished primarily by morphological features such as antennal structure, foreleg configuration, wing venation patterns, and male genital sclerites, which aid in taxonomic identification.^[39] Representative genera include Mantispa in Mantispinae and Symphrasis in Symphrasinae, reflecting the family's overall raptorial adaptations and ecological specializations.^[40] Recent taxonomic work has added new Neotropical species, enhancing understanding of regional diversity.^[19] Calomantispinae represents the basal extant subfamily, with about 10 species in 2 genera, primarily distributed in the Afrotropics and extending to the Neotropics.^[41] Members exhibit filiform antennae and simple spines on the forelegs, with wing venation showing reduced crossveins and relatively straight costal margins; male genitalia feature simple pseudopenes. Larvae are generalist predators targeting sedentary arthropods such as insect pupae, often boarding hosts directly.^[20] ^[40] Drepanicinae includes around 39 species across 5 genera, with a distribution centered in the Oriental and Australasian regions, and scattered occurrences in the Nearctic and Neotropics.^[42] Diagnostic traits encompass sickle-shaped raptorial forelegs adapted for grasping, filiform to slightly clavate antennae, and wing venation with prominent humeral veins and fewer branches in the radial sector; genital structures display complex parameres. Many species mimic wasps in coloration, and larvae are hypothesized to prey on lepidopterans or subterranean arthropods, though data remain limited.^[35] ^[20] Mantispinae is the largest and most diverse subfamily, encompassing approximately 320 species in 35 genera, with a global distribution but highest richness in the Neotropics.^[43] Key features include clubbed antennae, highly complex raptorial forelegs with multiple spine rows and articulated coxae, and wing venation characterized by forked media posterior veins and extensive reticulation; identification often relies on intricate male ectoproct and gonocoxite morphology. Larvae specialize in spider eggs, employing phoretic strategies to infiltrate sacs (see Larval Predation Strategies).^[44] ^[20] Symphrasinae contains about 36 species in 3 genera, exhibiting a pantropical distribution with extensions into the southern Nearctic.^[45] Distinguishing characteristics comprise an elongated prothorax, variable antennal forms from filiform to serrate, forelegs with moderate raptorial modifications, and wing venation featuring a bifurcated cubital vein and reduced anal branches; genital traits include distinctive hypandrium shapes. Forms range from ant mimics to soil-dwellers, with larvae employing diverse strategies such as targeting hymenopteran pupae in nests.^[20] ^[3]

Extinct Subfamilies and Fossil Record

The fossil record of Mantispidae extends back to the Early Jurassic, approximately 180 million years ago, with the earliest known specimens lacking fully developed raptorial forelegs, which first appear in the Middle Jurassic.^[21] The family experienced peak diversity during the Cretaceous, particularly in amber deposits, where over 50 extinct species have been documented, although this is fewer than the roughly 400 extant species today.^[46] Post-Cretaceous fossils are less common, with notable occurrences in Eocene and Oligocene compression deposits, and scattered Miocene amber inclusions, suggesting a decline in diversity after the Paleogene.^[47] Two subfamilies are exclusively extinct: †Mesomantispinae and †Doratomantispinae. The †Mesomantispinae, known from about 10 species primarily in the Early Cretaceous Yixian Formation of China, exhibit primitive wing venation patterns that position them as the basalmost lineage within Mantispidae.^[48] In contrast, †Doratomantispinae is restricted to mid-Cretaceous Kachin amber from Myanmar, with recent discoveries between 2020 and 2025 describing multiple new genera and highlighting their robust, raptorial forelegs adapted for predation.^[49] These subfamilies underscore the Mesozoic radiation of mantispids, with morphological disparity indicating greater ancient diversity than in modern faunas.^[38] Key fossil localities include the prolific mid-Cretaceous Myanmar (Kachin) amber, which has yielded numerous Doratomantispinae specimens and new genera described in 2023–2025, revealing fine details of raptorial limb evolution.^[49] Lebanese amber from the Early Cretaceous has preserved early mesomantispine-like forms, while the Eocene Green River Formation in North America produced Protonolima mantispinoformis, an adult mantispid with affinities to extant subfamilies.^[50] These sites demonstrate the global distribution of ancient mantispids and their adaptation to diverse paleoecosystems. Evolutionarily, the fossil record illustrates the early divergence of raptorial foreleg traits in Mantispidae, with Mesozoic forms showing varied spination and elongation that prefigure modern predatory specializations, likely driven by interactions with spider prey.^[51] This disparity, especially in Cretaceous amber, suggests higher lineage diversity in the past, possibly linked to broader ecological opportunities before a post-Paleogene contraction.^[48] Preservation in the fossil record favors adult stages, with over 80% of known specimens being winged imagines; larval fossils are exceedingly rare, limited to a handful of amber inclusions showing campodeiform first instars associated with spider egg sacs.^[52] No definitive post-Eocene larval fossils are known, further emphasizing the scarcity of immature stages. A notable recent discovery from 2025 includes a new genus of Drepanicinae from the earliest Eocene Fur Formation in Denmark, alongside Danomantispa and Protheristria, expanding the Paleogene record and highlighting northern high-latitude occurrences of mantispids.^[53]

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[43]
Symphrasinae - Explore the Taxonomic Tree | FWS.gov
Branches of Symphrasinae. Hide lower priority columns. Taxonomic Level, Taxa, FWS Focus. Tribe, 0. Subtribe, 0. Genus, 3. Subgenus, 0. Species, 36. Aberration ...
[44]
A camouflaged diversity: taxonomic revision of the thorny lacewing ...
Apr 30, 2024 · Species of the thorny lacewing subfamily Symphrasinae (Neuroptera: Rhachiberothidae) are revised. Prior to this work, 42 species were known ...
[45]
A remarkable new genus of Symphrasinae (Neuroptera
Sep 24, 2025 · A diverse fauna of Mantispidae has been recently discovered in mid-Cretaceous Kachin amber comprised of 29 species in 14 genera. Six genera ...
[46]
Mantispids in the fossil record - Dr. James E. Jepson
Jun 15, 2015 · The Lower Cretaceous mantispids are all from the subfamily Mesomantispinae, these are mostly known from the Yixian Formation of China, with many ...
[47]
Cretaceous diversity and disparity in a lacewing lineage of predators ...
Jun 3, 2020 · Mantidflies (Mantispidae) are an unusual and charismatic group of predatory lacewings (Neuroptera), whereby the adults represent a remarkable ...
[48]
New Cretaceous fossil mantispids highlight the palaeodiversity of ...
Aug 6, 2025 · This family is traditionally divided into two extinct subfamilies: Mesomantispinae and Doratomantispinae, and four extant subfamilies ...
[49]
A new genus of the mantispid-like Paraberothinae (Neuroptera
Aug 28, 2015 · The specialized probasitarsi in Paraberothinae, Rhachiberothinae (Berothidae) and Symphrasinae (Mantispidae) are considered as a homoiology ( ...
[50]
The associated evolution of raptorial foreleg and mantispid ...
Nov 2, 2023 · Among them, Mantispidae have an evolutionary history fluctuating over 200 million years, along with their raptorial forelegs having transformed ...Missing: traits | Show results with:traits
[51]
The ride of the parasite: a 100-million-year old mantis lacewing larva ...
Dec 22, 2018 · The grub-like morphology of the stage two and three larvae could most likely only evolve after the strategy of entering egg sacs had evolved as ...
[52]
Mantispidae) from the earliest Eocene Fur Formation, Denmark
The two genera known from the Fur Formation, Danomantispa Makarkin et al., 2025 and Protheristria gen. nov., are not closely related and ...