Mantispidae Explained

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

About 5mm47mm long and with a wingspan of 5-, 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.

Extant taxa based on Global Biodiversity Information Facility^[8] and extinct taxa based on Jepson, 2015 and subsequent literature.^[9]

Calomantispinae

• Calomantispa Banks, 1913
• Nolima Navás, 1914

Drepanicinae

• †Acanthomantispa Lu et al. 2020 Burmese amber, Myanmar, Late Cretaceous (Cenomanian)
• †Aragomantispa Pérez-de la Fuente and Peñalver 2019 Spanish amber, Early Cretaceous (Albian)^[10]
• †Dicranomantispa Lu et al. 2020 Burmese amber, Myanmar, Cenomanian
• Ditaxis (insect) McLachlan, 1867
• Drepanicus Blanchard, 1851
• Gerstaeckerella Enderlein, 1910
• †Liassochrysa Ansorge and Schlüter 1990 Green Series, Germany, Early Jurassic (Toarcian)
• †Promantispa Panfilov 1980 Karabastau Formation, Kazakshtan, Middle/Late Jurassic
• †Psilomantispa Lu et al. 2020 Burmese amber, Myanmar, Cenomanian
• †Sinuijumantispa So & Won, 2022 Sinuiju Formation, North Korea, Early Cretaceous (Aptian)
• Theristria Gerstaecker, 1884

Mantispinae

• Afromantispa Snyman & Ohl, 2012
• Asperala Lambkin, 1986
• Austroclimaciella Handschin, 1961
• Austromantispa Esben-Petersen, 1917
• Buyda (insect) Navás, 1926
• Campanacella Handschin, 1961
• Campion Navás, 1914
• Cercomantispa Handschin, 1959
• Climaciella Enderlein, 1910
• Dicromantispa Hoffman, 2002^[1]
• Entanoneura Enderlein, 1910
• Euclimacia Enderlein, 1910
• Eumantispa Okamoto, 1910
• †Feroseta Poinar 2006 Dominican amber, Miocene
• Haematomantispa Hoffman, 2002
• Leptomantispa Hoffman, 2002
• Madantispa Fraser, 1952
• Mantispa Illiger, 1798
• Mimetispa Handschin, 1961
• Nampista Navás, 1914
• Necyla Navás, 1913
• Nivella Navás, 1930
• Orientispa Poivre, 1984
• Paramantispa Williner & Kormilev, 1959
• Paulianella Handschin, 1960
• †Prosagittalata Nel 1988 Céreste, France, Rupelian
• Pseudoclimaciella Handschin, 1960
• Rectinerva Handschin, 1959
• Sagittalata Handschin, 1959
• Spaminta Lambkin, 1986
• Stenomantispa Stitz, 1913
• Toolida Lambkin, 1986
• Tuberonotha Handschin, 1961
• †Vectispa Lambkin 1986 Bembridge Marls, United Kingdom, Eocene (Priabonian)
• Xaviera (insect) Lambkin, 1986
• Xeromantispa Hoffman, 2002
• Zeugomantispa Hoffman, 2002

Symphrasinae

Auth: Navás, 1909

• †Archaeosymphrasis Shi et al. 2020 Burmese amber, Myanmar, Cenomanian
• Anchieta (insect) Navás, 1909
• †Habrosymphrasis Shi et al. 2020 Burmese amber, Myanmar, Cenomanian
• †Haplosymphrasites Lu et al. 2020 Burmese amber, Myanmar, Cenomanian
• †Parasymphrasites Lu et al. 2020 Burmese amber, Myanmar, Cenomanian
• Plega Navás, 1927 - Americas
• †Symphrasites Wedmann & Makarkin, 2007 Messel Pit, Germany, Eocene
• Trichoscelia Westwood, 1852

†Mesomantispinae

Auth: Makarkin 1996

• †Archaeodrepanicus Jepson et al. 2013 Yixian Formation, China, Early Cretaceous (Aptian)
• †Clavifemora Jepson et al. 2013 Daohugou, China, Middle/Late Jurassic
• †Karataumantispa Jepson 2015 Karabastau Formation, Kazakhstan, Middle/Late Jurassic
• †Mesomantispa Makarkin 1996 Zaza Formation, Russia, Aptian
• †Ovalofemora Jepson et al. 2018 Karabastau Formation, Kazakhstan, Middle/Late Jurassic
• †Sinomesomantispa Jepson et al. 2013 Yixian Formation, China, Aptian

Unassigned

• Allomantispa Liu, Wu, Winterton & Ohl, 2014
• Entatoneura Enderlein, 1910
• Fera (insect) Whalley, 1983
• Forciada Kozhanchikov, 1949
• Longicollum - monotypic Longicollum benmaddoxi Jepson et al., 2018
• Manega (insect) Navás, 1929
• †Neromantispa - monotypic Neromantispa antiqua Hart et al., 2024
• Promantispa Jarzembowski, 1980
• Prosagittalata Nel, 1988

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^[11] However, later studies found them to be basal members of Psychopsoidea and Neuroptera respectively.^[12]

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] [11]}

Notes and References

1. Engel . MS . Grimaldi . DA . 2007 . The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera) . American Museum Novitates . 3587 . 1–58 . 10.1206/0003-0082(2007)3587[1:TNFODA]2.0.CO;2. 2246/5880 . 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 . 10.1002/zoos.201000004.
4. Ohl . M . 2004 . A new wasp-mimicking species of the genus Euclimacia from Thailand (Neuroptera, Mantispidae) . Denisia . 13 . 193–196 .
5. Opler . PA . 1981 . Polymorphic Mimicry of Polistine Wasps by a Neotropical Neuropteran . Biotropica . 13 . 3 . 165–176 . 10.2307/2388121. 2388121 . 1981Biotr..13..165O .
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 . . 43 . 175–194 . 10.1146/annurev.ento.43.1.175. 15012388 .
8. https://www.gbif.org/species/5470 Global Biodiversity Information Facility: Mantispidae (retrieved 27 October 2020)
9. Jepson. James E.. 2015-06-04. A review of the current state of knowledge of fossil Mantispidae (Insecta: Neuroptera). Zootaxa. 3964. 4. 419–432. 10.11646/zootaxa.3964.4.2. 26249453. 1175-5334.
10. 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. en. 9. 1. 13248. 10.1038/s41598-019-49398-1. 2045-2322. 6744510. 31519980. 2019NatSR...913248P.
11. 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 . . 149 . 4 . 701–716 . 10.1111/j.1096-3642.2007.00273.x . free .
12. 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. en. 63. 1. 531–551. 10.1146/annurev-ento-020117-043127. 29324039 . 0066-4170.