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Comprehensive Description

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Around 400 species of mantispids, placed in around 44 genera, are known (Ohl and Oswald 2004). Diversity is higher at lower latitudes. For example, according to Reynoso-Velasco and Contreras-Ramos (2008), the known mantispid fauna of Mexico includes around two dozen species, whereas the (surely more complete) species list for all of the United States and Canada includes just half this number, nearly all of which are present in Mexico as well (and according to Cannings and Cannings [2006], only four species occur in Canada). Caribbean island diversity is equivalent to that of the entire Nearctic (Canada and the United States), although just a few species are not shared with continental areas. A summary of species richness by zoogeographic region reinforces this pattern, with the numbers of species known from the Nearctic or Palearctic regions being many times fewer than for other regions (Ohl and Oswald 2004), despite the fact that these two regions are presumably the most thoroughly known.

With the exception of the subfamily Mantispinae, little is known of the biology and ecology of mantispids. The Mantispinae, however, are relatively well known. In contrast to the scattered data available for the several other subfamilies, which suggest that a wide range of insects may be used as food by developing individuals of various mantispid species, known larval developmental associations of species in the Mantispinae are all with the egg sacs of spiders and the larvae are predators on spider eggs. Larvae of some mantispine species are obligate egg-sac penetrators. These larvae locate and penetrate spider egg sacs that have already been produced and deposited in the environment. Other mantispine species locate and board spiders, then enter the egg sac during its construction. Larvae of still other species use either or both techniques to get inside spider egg sacs. If larvae enter an egg sac containing nearly hatched spiderlings, they may board these. It is not clear whether larvae are able to search out spiders from a distance. Given that adults produce egg clutches containing from several hundred to several thousand eggs, it is possible that searching is purely random and that encounters with spiders are simply fortuitous. Although larvae would presumably benefit from boarding only female spiders, in all species that have been studied larvae board male and female spiders with equal frequency. However, in at least one species, larvae that find themselves on a male will transfer to the male's mate when he copulates; in at least one other species, larvae on a male spider will transfer to a female who cannibalizes the male. (Redborg 1998 and references therein)

Although it has long been known that mantispine larvae feed on spider eggs, only in the 1980s was it recognized that some (possibly all) spider boarders feed on spider hemolymph ("blood") while aboard adult spiders. Spider-boarding mantispids overwinter on their host spiders. The overwintering behavior of egg-sac penetrators is more varied. In some species, individuals overwinter as a mass of unfed first instars and search for egg sacs the following spring. Others may overwinter as larvae or pupae within spider egg sacs. (Redborg 1998 and references therein)

Once inside a spider egg sac, a mantispid larva pierces and drains eggs with its modified mandibles and maxillae. As a third instar larva, it constructs a pupal cocoon within the spider egg sac. In the laboratory at 25º C, the entire life cycle for the well studied Mantispa uhleri takes around 28 days from first instar to adult. The incipient adult bites its way out of both the cocoon and egg sac and walks some distance away before undergoing the final molt, presumably because egg sacs may be located in concealed or awkward places such as under tree bark or within a silken retreat. (Redborg 1998 and references therein)

Although data on which spider species are preferentially attacked are very incomplete (reviewed by Redborg 1998), some tentative generalizations are possible. Host range tends to be fairly broad, with few mantispine species believed to have a host range limited even to a single family. Egg sac-penetrating mantispids probably tend to have broader host ranges and to be more often associated with web-building spiders than are spider-boarding mantispids. Spider-boarding mantispids may be more likely to be associated with wandering spiders, while mantispids associated with the egg sacs of web-building spiders may be more likely to be egg-sac penetrators. At best, however, these are rough generalizations. (Redborg 1998 and references therein)

Little is known about the nonreproductive habits of adult mantispines. Several species appear to be wasp mimics. In the few mantispine species in which mating behavior has been observed, courtship begins with male and female facing each other. Both male and female slowly extend forward and flex the coxa and femur of one raptorial leg followed by identical movements of the other leg. Males possess a glandular abdominal epithelium that produces a suspected pheromone, although this requires further investigation. Copulation occurs venter to venter, which necessitates twisting of the male’s abdomen through a 180º angle to achieve the final orientation of male and female facing away from each other. Copulation duration ranges from 1 hour to 24 hours among species that have been studied. Postcopulatory sparring may occur. A white opalescent sperm-containing spermatophore, present at the tip of the female abdomen following mating, is generally absorbed within 24 to 48 hours. Mantispids lay discrete clutches of very large numbers of eggs, each of which is attached to the substratum by a short stalk produced by secretions from the female's colleterial ("glue") gland. Depending on the species and the size of an individual female, clutch size may vary from a couple of hundred to several thousand eggs, with one to two clutches produced per week for several weeks or even months. (Redborg 1998 and references therein)

Ohl and Oswald (2004) catalogued the world mantispid fauna, including all fossil and extant species known. They present data on original and current generic placements and synonymies and include literature sources, type status, type locality, type depository, and taxon distributions.

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Mantispidae

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Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide,[1] especially in the tropics and subtropics. Only five species of Mantispa occur in Europe.[2] As their names suggest, members of the group possess raptorial forelimbs similar to those of the praying mantis, a case of convergent evolution.

Description and ecology

Head of a mantisfly in genus Plega
Species unknown, Sydney, Australia
A Sagittalata species female from Kerala, India

About 5–47 mm (0.20–1.85 in) long and with a wingspan of 5–30 mm (0.2–1.2 in), some mantidflies such as Climaciella brunnea, Euclimacia nodosa[3][4] are wasp mimics,[5] but most are brownish with green, yellow and sometimes red hues. The vernacular and scientific names are derived from their mantis-like appearance, as their spiny "raptorial" front legs are modified to catch small insect prey and are very similar to the front legs of mantids (the only difference is that the pincers lack footpads and are not used for walking at all). The adults are predatory insects that are often nocturnal, and are sometimes attracted by porch lights or blacklights. They are usually green, brown, yellow, and sometimes pink, and have four membranous wings which may sometimes be patterned (especially in wasp mimicking species) but are usually clear. Adult mantidflies are predators of suitably sized insects, which they catch as mantids do. However, the underlying mechanisms for the prey capture behavior are different in mantidflies and mantids.[6] Mantidflies are active hunters, but as with other Neuroptera, they are cumbersome fliers.

Symphrasinae larvae are sedentary parasitoids on bee, wasp or scarab beetle larvae. Larvae of the Calomantispinae are predators of small arthropods, and in at least one species they are mobile. Mantispinae have the most specialized larval development among all mantidflies studied to date (the life history of the Drepanicinae remains unknown): their campodeiform larvae seek out female spiders or their egg sacs which they then enter; the scarabaeiform larvae then feed on the spider eggs, draining egg contents through a piercing/sucking tube formed by modified mandibles and maxillae, pupating in the egg sac.[1]

First-instar mantispids use two strategies to locate spider eggs: larvae may burrow directly through the silk of egg sacs they find, or they may board and be carried by female spiders prior to sac production (phoresy), entering the sac as it is being constructed. Mantispids that board spiders usually adopt positions on or near the pedicel; some species may enter the spider's book lungs. Larvae maintain themselves aboard spiders by feeding on spider hemolymph. Transfers of larvae from spider to spider are possible during spider mating or cannibalism. All of the major groups of hunting spiders are attacked by spider-boarding mantispids; the egg sacs of web-building species are also entered by egg-sac penetrators.[7]

Systematics

Among the Neuroptera (which includes lacewings and owlflies), mantidflies are apparently most closely related to the Dilaridae (pleasing lacewings) and the thorny (Rhachiberothidae) and beaded lacewings (Berothidae). These and the prehistoric Mesithonidae - probably a paraphyletic assemblage rather than a natural group - form the superfamily Mantispoidea.

Many mantidflies are placed in one of the four subfamilies, of which the Symphrasinae are probably the most distinct and the Mantispinae are the most advanced. But a considerable number of taxa cannot be easily accommodated in this layout, and are therefore better treated as incertae sedis at present.

Some authors have suggested that the extinct two winged Dipteromantispidae known from Cretaceous fossils should be treated as a subfamily of Mantispidae.[8]

Extant taxa based on Global Biodiversity Information Facility[9] and extinct taxa based on Jepson, 2015 and subsequent literature.[10]

Calomantispinae

Drepanicinae

Mantispinae

Symphrasinae

Auth: Navás, 1909

†Mesomantispinae

Auth: Makarkin 1996

Unassigned

Fossil taxa may be of an altogether quite basal position, for example the Jurassic Liassochrysa (about 180 million years old) and Promantispa (about 155 million years old) have been assigned to either a basal position within the group or Drepanicinae, the most basal subfamily within the group. The Early Jurassic Prohemerobius dilaroides (the type species of the "Prohemerobiidae" assemblage) as well as the Late Permian Permantispa emelyanovi (of the just as likely paraphyletic "Permithonidae") were suggested to possibly represent ancestral mantidflies[12] However, later studies found them to be basal members of Psychopsoidea and Neuroptera respectively.[8]

Most living genera from which fossil species are also known to go back to the Miocene; the Oligocene "Climaciella" henrotayi probably does not belong in the living genus. Two fossil species have been described as part of the extant genus Dicromantispa, Dicromantispa moronei from Dominican amber and Dicromantispa electromexicana from Mexican amber.[1]

The North American species include:

Paraberotha, Retinoberotha and Whalfera were formerly placed here, but have since been recognized as Rhachiberothidae. Mantispidiptera are diminutive insects, apparently neuropterans of some sort, perhaps Hemerobiiformia; their exact affiliation cannot at present be determined because of their odd apomorphies, though they are unlikely to have been mantidflies.[1][12]

References

  1. ^ a b c d e Engel, MS; Grimaldi, DA (2007). "The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera)". American Museum Novitates (3587): 1–58. doi:10.1206/0003-0082(2007)3587[1:TNFODA]2.0.CO;2. hdl:2246/5880. S2CID 49393365.
  2. ^ Aspöck, Ulrike & Aspöck, Horst (2010): Fauna Europaea – Mantispidae. Version of 2010-DEC-23. Retrieved 2011-JAN-03.
  3. ^ Bhattacharjee, S; Ohl, M; Saha, S; Sarkar, S; Raychaudhuri, D (2010). "Euclimacia nodosa (Westwood, 1847), a rare and poorly known species of Mantispidae (Neuroptera), recorded for the first time from West Bengal, India". Zoosystematics and Evolution. 86 (2): 221–224. doi:10.1002/zoos.201000004.
  4. ^ Ohl, M (2004). "A new wasp-mimicking species of the genus Euclimacia from Thailand (Neuroptera, Mantispidae)" (PDF). Denisia. 13: 193–196.
  5. ^ Opler, PA (1981). "Polymorphic Mimicry of Polistine Wasps by a Neotropical Neuropteran". Biotropica. 13 (3): 165–176. doi:10.2307/2388121. JSTOR 2388121.
  6. ^ Kral, K (2013). Vision in the mantispid: a sit-and-wait and stalking predatory insect. Physiological Entomology 38: 1-12.
  7. ^ Redborg, KE (1998). "Biology of the Mantispidae". Annual Review of Entomology. 43: 175–194. doi:10.1146/annurev.ento.43.1.175. PMID 15012388.
  8. ^ a b Engel, Michael S.; Winterton, Shaun L.; Breitkreuz, Laura C.V. (2018-01-07). "Phylogeny and Evolution of Neuropterida: Where Have Wings of Lace Taken Us?". Annual Review of Entomology. 63 (1): 531–551. doi:10.1146/annurev-ento-020117-043127. ISSN 0066-4170. PMID 29324039.
  9. ^ Global Biodiversity Information Facility: Mantispidae (retrieved 27 October 2020)
  10. ^ Jepson, James E. (2015-06-04). "A review of the current state of knowledge of fossil Mantispidae (Insecta: Neuroptera)". Zootaxa. 3964 (4): 419–432. doi:10.11646/zootaxa.3964.4.2. ISSN 1175-5334. PMID 26249453.
  11. ^ Pérez-de la Fuente, Ricardo; Peñalver, Enrique (2019-09-13). "A mantidfly in Cretaceous Spanish amber provides insights into the evolution of integumentary specialisations on the raptorial foreleg". Scientific Reports. 9 (1): 13248. Bibcode:2019NatSR...913248P. doi:10.1038/s41598-019-49398-1. ISSN 2045-2322. PMC 6744510. PMID 31519980.
  12. ^ a b Wedmann, S; Makarkin, VN (2007). "A new genus of Mantispidae (Insecta: Neuroptera) from the Eocene of Germany, with a review of the fossil record and palaeobiogeography of the family" (PDF). Zoological Journal of the Linnean Society. 149 (4): 701–716. doi:10.1111/j.1096-3642.2007.00273.x.

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Mantispidae: Brief Summary

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Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in the tropics and subtropics. Only five species of Mantispa occur in Europe. As their names suggest, members of the group possess raptorial forelimbs similar to those of the praying mantis, a case of convergent evolution.

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