A new genus of mantidflies discovered in the Oriental region,with a higher‐level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology

A remarkable new genus and two new species of Mantispidae (Neuroptera) are described from the Oriental region. Allomantispa Liu, Wu, Winterton & Ohl gen.n. , currently including A. tibetana Liu, Wu & Winterton sp.n. and A. mirimaculata Liu & Ohl sp.n. The new genus is placed in the subfamily Drepanicinae based on a series of morphological characteristics and on the results of total evidence phylogenetic analyses. Bayesian and Parsimony analyses were undertaken using three gene loci (CAD, 16S rDNA and COI) combined with 74 morphological characters from living and fossil exemplars of Mantispidae (17 genera), Rhachiberothidae (two genera) and Berothidae (five genera), with outgroup taxa from Dilaridae and Osmylidae. The resultant phylogeny presented here recovered a monophyletic Mantispidae with ?Mesomantispinae sister to the rest of the family. Relationships among Mantispidae, Rhachiberothidae and Berothidae support Rhachiberothidae as a separate family sister to Mantispidae. Within Mantispidae, Drepanicinae are a monophyletic clade sister to Calomantispinae and Mantispinae. In a combined analysis, Allomantispa gen.n. was recovered in a clade comprising Ditaxis McLachlan from Australia, and two fossil genera from the Palaearctic, ?Promantispa Panfilov (Kazakhstan; late Jurassic) and ?Liassochrysa Ansorge & Schlüter (Germany; Jurassic), suggesting a highly disjunct and relictual distribution for the family. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:464B06E8‐47E6‐482E‐8136‐83FE3B2E9D6B .     

A new genus of Mantispidae (Insecta: Neuroptera) from the Eocene of Germany, with a review of the fossil record and palaeobiogeography of the family

A new mantispid genus and species Symphrasites eocenicus gen. sp. nov. is described from the Middle Eocene of the Messel Pit fossil site, Germany. It has been placed, with confidence, in the subfamily Symphrasinae, the fossils of which were previously unknown. A review of the known fossil species of Mantispidae is provided, and their systematic placements are discussed. Liassochrysa stigmatica Ansorge & Schlüter, 1990 is assigned to Mantispidae, and therefore Liassochrysidae is a new synonym of Mantispidae; a second, more complete specimen of Mesomantispa sibirica Makarkin, 1997 is described from the Early Cretaceous Baissa locality, Russia; Mantispidiptera Grimaldi, 2000 is excluded from the family; Vectispa Lambkin, 1986 is referred to subfamily Mantispinae; and the assignment of Whalfera Engel, 2004 to Rhachiberothidae is validated. The origin and palaeobiogeography of the mantispids are briefly discussed: the concept of ousted relicts – which assumes that former widely distributed taxa were replaced with groups originating in the tropics – is assumed to best explain the present distribution of the family.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 701–716.     

Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera)

A phylogenetic analysis of Neuroptera using thirty‐six predominantly morphological characters of adults and larvae is presented. This is the first computerized cladistic analysis at the ordinal level. It included nineteen species representing seventeen families of Neuroptera, three species representing two families (Sialidae and both subfamilies of Corydalidae) of Megaloptera, two species representing two families of Raphidioptera and as prime outgroup one species of a family of Coleoptera. Ten equally most parsimonious cladograms were found, of which one is selected and presented in detail. The results are discussed in light of recent results from mental phylogenetic cladograms. The suborders Nevrorthi‐ formia, Myrmeleontiformia and Hemerobiiformia received strong support, however Nevrorthiformia formed the adelphotaxon of Myrmeleontiformia + Hemerobiiformia (former sister group of Myrmeleontiformia only). In Myrmeleontiformia, the sister‐group relationships between Psychopsidae + Nemopteridae and Nymphidae + (Myrmeleontidae + Ascalaphidae) are corroborated. In Hemerobiiformia, Ithonidae + Polystoechotidae is confirmed as the sister group of the remaining families. Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)), which has already been proposed, is confirmed. Chrysopidae + Osmylidae emerged as the sister group of a clade comprising Hemerobiidae + ((Coniopterygidae + Sisyridae) + (dilarid clade)). Despite the sister‐group relationship of Coniopterygidae + Sisyridae being only weakly supported, the position of Coniopterygidae within the higher Hemerobiiformia is corroborated. At the ordinal level, the analysis provided clear support for the hypothesis that Megaloptera + Neuroptera are sister groups, which upsets the conventional Megaloptera + Raphidioptera hypothesis.     

Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola)

Abstract Segment 9 of male Raphidioptera, comprising tergite, sternite, gonocoxites, gonostyli and gonapophyses, is a benchmark for homologies in the male and female terminalia of the three Neuropterida orders Raphidioptera, Megaloptera and Neuroptera. The segments relating to genitalia are 9, 10 and 11 in males and 7, 8 and 9 in females. Results from holomorphological and recent molecular cladistic analyses of Neuropterida agree in supporting the sister‐group relationships between: (1) the Raphidioptera and the clade Megaloptera + Neuroptera, and (2) the suborder Nevrorthiformia and all other Neuroptera. The main discrepancy between the results of these studies is the nonmonophyly of the suborder Hemerobiiformia in the molecular analysis. The monophyly of the Megaloptera (which has been repeatedly questioned) is further corroborated by a hitherto overlooked ground pattern autapomorphy: the presence of eversible sacs within the complex of the fused gonocoxites 11 in Corydalidae and Sialidae. The recently discovered paired complex of gonocoxites 10 (parameres) in Nipponeurorthus (Nevrorthidae) indicates that the curious apex of sternite 9 of Nevrorthus and Austroneurorthus is the amalgamation of the sclerites of gonocoxites 10 with sternite 9, interpreted as synapomorphic. In the molecular study, the Nevrorthidae, Sisyridae and Osmylidae branch off in consecutive splitting events, a result that is supported by the analysis of male genital sclerites reported here. Extraordinary parallel apomorphies (e.g. excessive enlargement and modification of gonocoxites 10 ending in a thread‐like ‘penisfilum’) in derived representatives of Coniopterygidae, Berothidae, Rhachiberothidae and Mantispidae corroborate the dilarid clade of the morphological analysis and leads us to hypothesize a sister‐group relationship of the Coniopterygidae with the dilarid clade. A re‐interpretation of the tignum of Chrysopidae as gonocoxites 11 means that the structure previously called the gonarcus represents the fused gonocoxites 9. In Hemerobiidae, the corresponding sclerite is consequently also homologized as fused gonocoxites 9. The enlargement of the lateral wings of the gonocoxites in both families is interpreted as a synapomorphy. Excessive enlargement of gonostyli 11 in the Polystoechotid clade and Myrmeleontiformia supports a sister‐group relationship of these two clades. The occurrence of certain serial homologues of female genitalia structures (gonocoxites and gonapophyses), such as the digitiform processus together with the flat appendices in segment 8 of certain Myrmeleontidae, or the wart‐like processus together with the flat circular sclerites in segment 7 of certain Berothidae, as well as the presence of gonocoxites 8 as pseudosternites in certain Nemopteridae and Coniopterygidae, are probably character reversals. The digitiform processus of tergite 9 (pseudogonocoxites) in Rhachiberothidae and Austroberothella (Berothidae) are either independently developed acquisitions with a function in oviposition, or are homologous sclerites, possibly of epipleurite origin.     

Two new species of Sinosmylites Hong (Neuroptera, Berothidae) from the Middle Jurassic of China, with notes on Mesoberothidae

Two new species of the genus Sinosmylites Hong are described from the Middle Jurassic locality at Daohugou (Inner Mongolia, China): Sinosmylites fumosussp. n. and Sinosmylites rasnitsynisp. n. This is the oldest known occurrence of the family Berothidae. The berothid affinity of this genus is confirmed by examination of the hind wing venation characteristic of the family. The Late Triassic family Mesoberothidae may represent an early group of Berothidae.     

The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta)

External and internal head structures of larvae of Nevrorthidae were described in detail. The results were compared to conditions found in other representatives of Neuroptera and the other two neuropterid orders. The cladistic analysis supported the monophyly of Neuroptera, Neuroptera exclusive of Nevrorthidae, Hemerobiiformia, and Myrmeleontiformia. Neuroptera exclusive of Nevrorthidae are supported by the formation of an undivided postmentum and the presence of cryptonephric Malpighian tubules. The highly specialized articulation of the neck (Rollengelenk) and the absence of a salivary duct are autapomorphies of Nevrorthidae. Ithonidae and Polystoechotidae form a clade and are the sister group of the remaining Hemerobiiformia, which are characterized by the complete lack of a gula and a terminal filament of the antenna. Within this lineage, a clade comprising Mantispidae, Dilaridae, Berothidae, and Rhachiberothidae is well supported. Larvae of Myrmeleontiformia are characterized by a complex transformation of head structures, with a hypostomal bridge, a small triangular gula, largely reduced maxillary grooves, and anteriorly shifted posterior tentorial grooves. The slender finger‐like mid‐dorsal apodeme is another autapomorphy of the group. Psychopsidae are placed as the sister group of the remaining Myrmeleontiformia, which are characterized by a conspicuous, protruding ocular region (often less distinct or even absent in Nemopteridae). Ascalaphidae are the sister group of Myrmeleontidae. Larvae of both families share the fusion of the tibia and tarsus in the hind leg. The larval characters analysed were not sufficient for full resolution of the myrmeleontiform and hemerobiiform lineages. The position of several families such as Osmylidae, Sisyridae, and Coniopterygidae remains uncertain. The results are in agreement with an aquatic ancestor of Neuroptera and secondarily acquired terrestrial habits within the lineage (Neuroptera exclusive of Nevrorthidae), and another invasion of the aquatic environment by Sisyridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 533–562.     

A revision and key to the genera of Afrotropical Mantispidae (Neuropterida, Neuroptera), with the description of a new genus

The Afrotropical Mantispidae genera have previously been neglected and are poorly known. The genera are revised and redescribed. A new genus Afromantispa Snyman and Ohl is described with Afromantispa tenellacomb. n.as type species. Perlamantispa (Handschin, 1960) is synonymised with Sagittalata Handschin, 1959. The new combinations within the genus include Sagittalata austroafricacomb. n., Sagittalata bequaerticomb. n., Sagittalata dorsaliscomb. n., Sagittalata girardicomb. n., Sagittalata nubilacomb. n.,Sagittalata perlacomb. n.,Sagittalata pusillacomb. n., Sagittalata similatacomb. n., Sagittalata royicomb. n., Sagittalata tinctacomb. n. andSagittalata vasseicomb. n. An illustrated key to the genera Afromantispagen. n., Sagittalata Handschin, 1959, Mantispa Illiger, 1798, Cercomantispa Handschin, 1959, Rectinerva Handschin, 1959, Nampista Navás, 1914, and Pseudoclimaciella Handschin, 1960 is provided. The wing venation of Mantispidae is redescribed. Similarities between the genera are discussed. Subsequent studies will focus on revising the taxonomic status of species, which are not dealt with in this study.     

Ultrastructural studies on euspermatozoa and paraspermatozoa in Mantispidae (Insecta, Neuroptera)

Previous studies have demonstrated the presence of sperm dimorphism in the Mantispidae Perlamantispa perla. We extended the study on several other mantidflies. In all the examined species the occurrence of euspermatozoa (typical) and paraspermatozoa (atypical) was established. The euspermatozoa are characterized by the presence of a cylindrical nucleus surrounded by an envelope that fans out laterally into two thin wings of different length. The acrosome seems to be missing. The nucleus is surrounded by extracellular material. The flagellum is provided with a 9 + 9 + 2 axonemal pattern; the accessory tubules contain 16 protofilaments and the intertubular material has the distribution typical of the taxon. Two elongated accessory bodies flank partially the axoneme and connect this structure with the mitochondrial derivatives. The flagellar axoneme of paraspermatozoa consists of an axoneme and two giant mitochondrial derivatives filled with large globular units. The axoneme exhibits a 9 + 9 + 2 pattern, in which the central 9 + 2 units have a normal structure, in that the microtubular doublets are provided with both dynein arms and radial links. On the contrary, the nine accessory microtubules have a large diameter and their tubular wall consists of 40 protofilaments. This comparative study provided evidences about the uniformity of sperm ultrastructure in Mantispidae. The function of non-fertilizing giant sperm in mantidflies is discussed.     

New fossil mantidflies (Insecta: Neuroptera: Mantispidae) from the Mesozoic of north‐eastern China

Abstract:  Three new genera and four new species of the extinct mantidfly subfamily Mesomantispinae (Insecta: Neuroptera: Mantispidae) are described from the Lower Cretaceous Yixian Formation of Liaoning and the Middle Jurassic Jiulongshan Formation of Inner Mongolia: Archaeodrepanicus nuddsi gen. et sp. nov., A. acutus gen. et sp. nov., Sinomesomantispa microdentata gen. et sp. nov., (Yixian Formation) and Clavifemora rotundata gen. et sp. nov. (Jiulongshan Formation). The specimens described herein represent the first Mesozoic mantidfly compression fossils to have body parts preserved, including the specialized raptorial forelegs articulated to the prothorax anteriorly, an autapomorphy of the family. These new taxa further confirm the placement of the subfamily Mesomantispinae within the family Mantispidae; however, the monophyly of Mesomantispinae has not been confirmed, and it is likely that it will prove to be paraphyletic.     

A new species of Stenobiella Tillyard (Neuroptera, Berothidae) from Australia

Stenobiella variolasp. n., a new species of beaded lacewing (Neuroptera: Berothidae), is described and figured from south-eastern Australia. A preliminary key to Stenobiella species is presented.     

Nocturnal flight activity of some African Mantispidae (Neuroptera)

An account is given of catches of six species of Mantispidae in a light-trap at Entebbe, Uganda, in 1961–62.     

Phylogeny of the Neuropterida (Insecta: Holometabola)

The Neuropterida, with about 6500 known species — living fossils in a way — at the base of the Holometabola (as a sister group of the Coleoptera), comprise Raphidioptera (about 210 species, two families), Megaloptera (about 300 species, two families) and Neuroptera (6000 species, 17 families). Megaloptera + Neuroptera is argued vs. the traditional Raphidioptera + Megaloptera. Raphidioptera are undisputedly monophyletic. Monophyly of Megaloptera is the operational hypothesis, although occasionally questioned. Sucking tubes of the larvae are the most spectacular autapomorphy of Neuroptera. The construction of larval head capsules indicates three evolutionary lines: Nevrorthiformia, and Myrmeleontiformia + Hemerobiiformia. Traditional Myrmeleontiformia is Psychopsidae + (Nemopteridae + (Nymphidae + (Myrmeleontidae + Ascalaphidae))), the present approach is (Psychopsidae + Nemopteridae) + all other Myrmeleontiformia. Hemerobiiformia are based on the ‘maxillary head’ concept. The ithonid clade Ithonidae/Rapismatidae + Polystoechothidae and the dilarid clade Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)) are based on robust criteria. Other relationships remain unclear: Hemerobiidae + Chrysopidae (on similarity) and the ‘early offshoot’ concept of coniopterygidae (on autapomorphies) should not be perpetuated. Chysopidae + Osmylidae and (Hemerobiidae + (Coniopterygidae + Sisyridae)) + dilarid clade are discussed. Aquatic larvae, regarded as independent apomorphies of megaloptera and neuropteran Nevrorthidae and Sisyridae for a long time, are re‐interpreted as a synapomorphy of Megaloptera + Neuroptera and thus plesiomorphic within these groups. Terrestrial larvae (with cryptonephry to solve osmotic problems) are consequently apomorphic. Aquatic Sisyridae with cryptonephry of a single malpighian tubule, is conflicting, but larvae may have become secondarily aquatic, after a terrestrial intermezzo.     

Larvae of the Japanese termitophilous predator Isoscelipteron okamotonis (Neuroptera,Berothidae) use their mandibles and silk web to prey on termites

Several species of Berothidae are known to be specialist predators of termites during their larval stages. A female of the Japanese berothid Isoscelipteron okamotonis (Nakahara, 1914) was captured in the field and laid eggs in captivity. Of 67 eggs, 20 hatched into larvae, 3 of 9 which pupated and 1 of which reached the adult stage. The first and third instars preyed on workers of the termite Reticulitermes speratus, although the second instars did not. Larvae did not show the specific predatory behaviors observed in Northern American relatives. Instead, they bound paralyzed termites with silk that was apparently used for hunting. This is the first record of an Old World berothid raised from egg to adult in the laboratory.     

Doratomantispa burmanica n. gen., n. sp. (Neuroptera: Mantispidae), a new genus of mantidflies in Burmese amber

A new genus and new species of mantidflies, Doratomantispa burmanica n. gen., n. sp. (Neuroptera: Mantispidae), is described from Burmese amber. Diagnostic characters of the new genus are small body size, trichosors present around entire wing margin except basally, protarsus 5-segmented with paired, simple claws but no aroleum, profemur bearing six cuticular spines, inner surface of protibia with row of peg-like protrusions, Sc meets R₁ in region of pterostigma, costal space greatly narrows toward wing apex, with 16 veinlets in costal space on front wing while costal veinlets on hind wing are replaced by trichosors and CuP absent in hind wing. The abdomen of the mantidfly is filled with large spheres resulting from a possible rickettsial infection. Phoretic heterostigmatid mites are adjacent to the wings of the fossil.     

Interference by the mantispid Mantispa uhleri with the development of the spider Lycosa rabida

Abstract. 1. The larvae of Mantispa uhleri Banks (Neuroptera: Mantispidae) board spiders to await the production of an egg sac containing their obligate developmental food. While aboard the spider, larvae maintain themselves by feeding on spider blood. This parasitic behaviour was investigated by allowing larvae to board sixth instar Lycosa rabida Walckenaer (Araneae: Lycosidae). Larval parasitism has a direct and indirect effect on the developmental physiology of the spider.
2. The direct effect, equal in both spider sexes, is an increase in development time and a decrease in adult size.
3. The indirect effect on development time and adult size is brought about by the loss of an instar in female spiders only. Parasitized females were mature at nine or ten instars; control females at ten or eleven. Male instar number was not affected; both control and parasitized males were mature at nine or ten instars.
4. The net result is that parasitized female spiders are even smaller than would be predicted from the direct effect alone, but actually mature faster than control females. In males there is only the direct effect. The adaptive significance of this sexually dimorphic response is discussed.     

New records of rare insects in western Transbaikalia

Marumba gaschkewitschii Bremer et Grey, 1852 (Lepidoptera, Sphingidae) is found in western Transbaikalia for the first time. Populations of the following rare insects previously known in the region from solitary specimens are found and described: Niphanda fusca Bremer et Grey, 1853, Plebejidea cyane Eversmann, 1837 (Lepidoptera, Lycaenidae), and Mantispa lobata Navás, 1912 (Neuroptera, Mantispidae). These populations are associated with climatic and microclimatic conditions of dense shrubs with almond trees.     

Vision in the mantispid: a sit‐and‐wait and stalking predatory insect

Mantispids (Neuroptera: Mantispidae) are remarkable insects as a result of their close resemblance to the praying mantis (order Mantodea). Although not closely related phylogenetically, as a result of similar selective pressures, both mantispids and mantids have evolved powerful raptorial forelegs for capturing insects. Another striking feature is the hypermetamorphosis in mantispid development, as well as the parasitizing behaviour of the first‐instar larvae. The present review focuses on the role of mantispid vision. First, the morphology and functional significance of the larval eyes (stemmata) are examined. In principle, the stemmata are suitable for spatial vision because of their arrangement and structure. This is then followed by a discussion of how adult mantispids are able to capture fast‐moving insects successfully, although, in contrast to the praying mantis, mantispids rely on superposition eyes rather than on apposition eyes with a frontal region of high acuity. For both larvae and adults, comparisons are made with other insect groups. The present review also addresses the role of mantispid vision as an important cue for triggering mating behaviour; accordingly, sex‐specific differences are considered. Finally, vision in the context of orientation flight is discussed.     

Protoresinacarus brevipedis gen. n., sp. n. from Early Cretaceous Burmese amber: the first fossil record of mites of the Family Resinacaridae (Acari: Heterostigmata: Pyemotoidea)

A new genus and species of mites, Protoresinacarus brevipedis gen. n., sp. n. (Acari: Heterostigmata: Pyemotoidea), is described from Early Cretaceous Burmese amber. This represents the first fossil record of a member of the family Resinacaridae. It is represented by 21 phoretic females adjacent to an adult mantidfly (Neuroptera: Mantispidae). This is the first record of phoresy of pyemotid mites on members of the insect order Neuroptera. The fossil mites differ from extant members of the family in possessing distinctly shorter legs I, which do not reach beyond the apex of the gnathosoma, and by the long setae v ₁, v ₂ and c ₂.