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Mantispidae (), commonly known as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of small to moderate-sized
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s in the order
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
. There are many
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
with around 400
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
worldwide, especially in the
tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
subtropics The subtropical zones or subtropics are geographical and climate zones immediately to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately ...
. Only five species of '' Mantispa'' occur in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. As their names suggest, members of the group possess
raptorial In biology (specifically the anatomy of arthropods), the term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod leg, arthropod's foreleg that make it function for the grasping of prey whi ...
forelimbs similar to those of the
praying mantis Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate ...
, a case of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
.


Description and ecology

About long and with a wingspan of , some mantidflies such as '' Climaciella brunnea'', '' Euclimacia nodosa'' are
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
mimic 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. In the simples ...
s, but most are brownish with green, yellow and sometimes red hues. The vernacular and scientific names are derived from their
mantis Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate a ...
-like appearance, as their spiny "
raptorial In biology (specifically the anatomy of arthropods), the term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod leg, arthropod's foreleg that make it function for the grasping of prey whi ...
" 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
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
y insects that are often
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, and are sometimes attracted by porch lights or
blacklight A blacklight, also called a UV-A light, Wood's lamp, or ultraviolet light, is a lamp that emits long-wave ( UV-A) ultraviolet light and very little visible light. One type of lamp has a violet filter material, either on the bulb or in a se ...
s. 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
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s of suitably sized
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s, which they catch as mantids do. However, the underlying mechanisms for the prey capture behavior are different in mantidflies and mantids. Mantidflies are active hunters, but as with other Neuroptera, they are cumbersome fliers. Symphrasinae larvae are sedentary
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s on bee,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
or scarab beetle larvae. Larvae of the Calomantispinae are
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s of small
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, 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
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s 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. 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 Phoresis or phoresy is a temporary commensalistic relationship when an organism (a phoront or phoretic) attaches itself to a host organism solely for travel. It has been seen in ticks and mites since the 18th century, and in fossils 320  ...
), 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 lung A book lung is a type of respiration organ used for atmospheric gas-exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open, ventral-abdominal, air-filled cavity (atrium) and co ...
s. Larvae maintain themselves aboard spiders by feeding on spider
hemolymph Hemolymph, or haemolymph, is a fluid, similar to the blood in invertebrates, that circulates in the inside of the arthropod's body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph c ...
. 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.


Systematics

Among the
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
(which includes
lacewing The Hemerobiiformia are a suborder of insects in the order Neuroptera that include most of the lacewings, antlions and their allies. The phylogeny of the Neuroptera was explored in 2014 using mitochondrial DNA sequences. The results indicate t ...
s, antlions 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 Prehistory, also called pre-literary history, is the period of human history between the first known use of stone tools by hominins  million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
Mesithonidae - probably a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
assemblage rather than a natural group - form the superfamily Mantispoidea. Many mantidflies are placed in one of the four
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
, of which the Symphrasinae are probably the most distinct and the Mantispinae are the most advanced. But a considerable number of
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
cannot be easily accommodated in this layout, and are therefore better treated as ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' 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 The Global Biodiversity Information Facility (GBIF) is an international organisation that focuses on making scientific data on biodiversity available via the Internet using web services. The data are provided by many institutions from around th ...
Global Biodiversity Information Facility: Mantispidae (retrieved 27 October 2020)
/ref> and extinct taxa based on Jepson, 2015 and subsequent literature.


Calomantispinae

* '' Calomantispa'' Banks, 1913 * '' Nolima'' Navás, 1914


Drepanicinae

* †'' Acanthomantispa'' Lu et al. 2020
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...
, Myanmar, Late Cretaceous (
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
) *†'' Aragomantispa'' Pérez-de la Fuente and Peñalver 2019 Spanish amber, Early Cretaceous (
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
) *†'' Dicranomantispa'' Lu et al. 2020 Burmese amber, Myanmar, Cenomanian *'' Ditaxis'' McLachlan, 1867 * '' Drepanicus'' Blanchard, 1851 * '' Gerstaeckerella'' Enderlein, 1910 *†'' Liassochrysa'' Ansorge and Schlüter 1990 Green Series, Germany, Early Jurassic (
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
) *†'' 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 The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
) * '' Theristria'' Gerstaecker, 1884


Mantispinae

* '' Afromantispa'' Snyman & Ohl, 2012 * '' Asperala'' Lambkin, 1986 * '' Austroclimaciella'' Handschin, 1961 * '' Austromantispa'' Esben-Petersen, 1917 * '' Buyda'' Navás, 1926 * '' Campanacella'' Handschin, 1961 * '' Campion'' Navás, 1914 * '' Cercomantispa'' Handschin, 1959 * '' Climaciella'' Enderlein, 1910 * '' Dicromantispa'' Hoffman, 2002 * '' Entanoneura'' Enderlein, 1910 * '' Euclimacia'' Enderlein, 1910 * '' Eumantispa'' Okamoto, 1910 *†'' Feroseta'' Poinar 2006
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil in ...
,
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
* '' 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 The Rupelian is, in the geologic timescale, the older of two age (geology), ages or the lower of two stage (stratigraphy), stages of the Oligocene epoch (geology), Epoch/series (stratigraphy), Series. It spans the time between . It is preceded b ...
* '' 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 The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(
Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage ...
) * '' Xaviera'' Lambkin, 1986 * '' Xeromantispa'' Hoffman, 2002 * '' Zeugomantispa'' Hoffman, 2002


Symphrasinae

Auth: Navás, 1909 * †'' Archaeosymphrasis'' Shi et al. 2020 Burmese amber, Myanmar, Cenomanian * '' Anchieta'' 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 The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.''Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sing ...
* †'' Symphrasites'' Wedmann & Makarkin, 2007
Messel Pit The Messel Formation is a Formation (geology), geologic formation in Hesse, central Germany, dating back to the Eocene Epoch (geology), epoch (about 47 Ma). Its geographic range is restricted to the Messel pit. There it unconformably overlie ...
, Germany,
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
*'' Trichoscelia'' Westwood, 1852


†Mesomantispinae

Auth: Makarkin 1996 * †'' Archaeodrepanicus'' Jepson et al. 2013
Yixian Formation The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
, China, Early Cretaceous (
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
) * †'' Clavifemora'' Jepson et al. 2013 Daohugou, China, Middle/Late Jurassic * †'' Karataumantispa'' Jepson 2015 Karabastau Formation, Kazakhstan, Middle/Late Jurassic * †'' Mesomantispa'' Makarkin 1996
Zaza Formation The Zaza Formation is a geological formation located in Buryatia (Russia). It dates to the Lower Cretaceous. The age of the formation is disputed, and is considered likely to be Valanginian-Hauterivian, or Aptian in age. It comprises sandstones, ...
, 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'' Whalley, 1983 * '' Forciada'' Kozhanchikov, 1949 * ''Longicollum'' - monotypic '' Longicollum benmaddoxi'' Jepson et al., 2018 * '' Manega'' Navás, 1929 * †''Neromantispa'' - monotypic '' Neromantispa antiqua'' Hart et al., 2024 * '' Promantispa'' Jarzembowski, 1980 * '' Prosagittalata'' Nel, 1988
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 preserve ...
taxa may be of an altogether quite basal position, for example the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
'' 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 The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...
'' Prohemerobius dilaroides'' (the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the " Prohemerobiidae" assemblage) as well as the
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
'' Permantispa emelyanovi'' (of the just as likely
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
" Permithonidae") were suggested to possibly represent ancestral mantidflies. However, later studies found them to be basal members of Psychopsoidea and
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
respectively. Most living genera from which fossil species are also known to go back to the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
; the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
''"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 Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil in ...
and '' Dicromantispa electromexicana'' from Mexican amber. 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
neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
ns of some sort, perhaps Hemerobiiformia; their exact affiliation cannot at present be determined because of their odd
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...
ies, though they are unlikely to have been mantidflies.


References


External links

{{Authority control Mantispoidea Insects used as insect pest control agents Neuroptera families Extant Early Jurassic first appearances