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Larval stages of Osmylidae: A. Kempynus sp. (photo: Kristi Ellington); B. Isostenosmylus sp. (photo: Enio Branco); C. unidentified stenosmyline (photo: Kristi Ellington).
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The genera of the lance lacewing family Osmylidae (Neuroptera) and extinct allied families (Archeosmylidae, Saucrosmylidae) are reviewed. A key to living Osmylidae genera of the world is presented. Each genus is diagnosed and figured with a checklist of species included for each. New, revised or support for previous taxonomic changes include the fo...
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... et al., 2018). Adults of all three families range in size from medium to relatively large (wing span ranging from 15 mm to over 120 mm), often with broad, distinctively patterned wings (e.g., Figs. 25, 48). Larvae of Osmylidae are campodeiform in shape with dramatically elongate lance-like jaws that are typically much longer than the head length (Fig. 1). The larvae are generalist predators either in leaf litter and under bark in some subfamilies (e.g., Stenosmylinae, Porisminae), or in riparian lotic habitats where they are often found under rocks and amongst debris close to the water edge (e.g., Kempyninae, Osmylinae) ( Winterton et al., 2017;Martins et al., 2018). Osmylidae, ...
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... so, relatively short; pterostigma very long and narrow; CuA with at least six pectinate branches. Comments. Members of Archeosmylus are relatively small species (wing length < 10 mm) with simplified venation and few medial vein branches. The absence of gradates is diagnostic for the genus although they are present in A. costalis (see Riek, 1955: fig. 21). The latter species is represented by an incomplete forewing and may belong to other genus and even ...
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... lobe present; female gonocoxite 9 (gx 9) (i.e., gonopophysis lateralis) elongate and with terminal stylus; female with a pair of sclerotised spermathecae; male genitalia with well-developed arched gonarcus with paired entoprocesses fused laterally; mediuncus curved; parameres present or absent, sometimes fused into single arched sclerite; larva ( Fig. 1) with straight, highly elongate jaws; gular-like sclerite absent; seven Malphigian tubules, of which five are incorporated into the cryptonephridium; eversible claws present on paired prolegs on last abdominal ...
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... 9, 29, 42); female sternite 8 reduced to small knob-like sclerite located immediately anterior to the lower margin of tergite 9, distant from sternite 7 (note: female genitalia unknown in Gumillinae HW area between MA-MP usually distinctly expanded, MP often forking in distal third (sometimes absent), branches of MP bifurcate near wing margin (Fig. 16); wing frequently falcate; male genitalia with parameres absent; female genitalia with gonapophysis 9 enlarged located anterior to ventral portion of tergite 9, often ornately modified and lobed (southern hemisphere (Fig. 4); wing never falcate; male genitalia with parameres present as paired rod-like structures lateral to mediuncus; ...
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... MA-MP fork relatively wide, crossveins between branches somewhat sinuous; large species (Australia, New Zealand and western South America) (Figs 13-16 Wings distinctly falcate; small species (forewing length <15.0 mm) (New Zealand) (Fig. 12) (Fig. 10) . . . . .Australysmus Kimmins. ...
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... MA-MP fork relatively wide, crossveins between branches somewhat sinuous; large species (Australia, New Zealand and western South America) (Figs 13-16 Wings distinctly falcate; small species (forewing length <15.0 mm) (New Zealand) (Fig. 12) (Fig. 10) . . . . .Australysmus Kimmins. ...
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... MA-MP fork relatively wide, crossveins between branches somewhat sinuous; large species (Australia, New Zealand and western South America) (Figs 13-16 Wings distinctly falcate; small species (forewing length <15.0 mm) (New Zealand) (Fig. 12) (Fig. 10) . . . . .Australysmus Kimmins. ...
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... species. C. sibiricus Makarkin, 1990 (Russia) (Early Cretaceous) Euosmylus Krüger, 1913a (Fig. 12) Type species. Stenosmylus stellae McLachlan, 1899: 259 (by monotypy) (New ...
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... different species and maybe even subfamilies. Lambkin (1987) mentioned some differences between the two but consider them as examples of intraspecific variation. The hind wing venation of the holotype is similar to Euporismus (Stenosmylinae) as believed by Tillyard & Dunstan (1916) and Riek (1952) (cf. Tillyard, 1916, fig. 1, and Lambkin, 1987 fig. 1). The incomplete preservation of the holotype does not allow precise determination of the subfamily, but the venation of the Dinmore specimen certainly belongs to Kempyninae based on the presence of the basal sinuous crossvein 1r-m and the deeply forked MP. E. balli Tillyard in Tillyard & Dunstan, 1916 (Australia Diagnosis. Forewing ...
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... that acts as a receptacle for gonapophysis 9; gonapophysis 9 enlarged, complex, variously bilobed, located anteriorly to ventral part of tergite 9, not closely associated with gonocoxite 9; gonocoxite 9 often with dark longitudinal stripe; spermatheca ovoid to spherical. Larvae typically have a pale to cream colouration with light brown sclerites (Fig. 1B, ...
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Citations
... The diversity, abundance and distribution of Neuroptera in forests can be affected by habitat types and forms, and the altitude of these environments can influence the fauna of some families (Bozdoğan 2020;Cancino-López et al. 2022). Osmylids can be found in humid habitats close to aquatic ecosystems (e.g., Osmylinae and Kemypninae) or living in leaf litter and under bark (e.g., Eidoporisminae, Porisminae, and Stenosmylinae) (New 1974;Cover and Resh 2008;Winterton et al. 2010Winterton et al. , 2019. ...
... Osmylidae (Neuroptera), commonly known as lance lacewings, are present in almost all biogeographical regions, except for the Nearctic, where fossil records indicate that the family has recently gone extinct there (Oswald 1994;Shepard and Contreras-Ramos 2009;Ardila-Camacho and Noriega 2014;Martins et al. 2016;Winterton and Wang 2016;Winterton et al. 2019). According to Shepard and Contreras-Ramos (2009), Osmylidae and other aquatic Neuroptera have no economic importance, not even as bioindicators, nevertheless, this still needs to be confirmed with further studies. ...
... There are about 280 described species of Osmylidae distributed in 25 living and 38 extinct genera (Winterton et al. 2019 Navás, 1912 (Gumillinae) are known from Brazil Martins et al. 2016Martins et al. , 2019Machado and Martins 2022), all of them from the Atlantic rainforest biome in the South and Southeast regions of the country (Machado and Martins 2022). As large territorial extensions of the country remain little explored, the fauna of Osmylidae in many Brazilian biomes is certainly underrepresented, for example, a new species of Isostenosmylus is being described to northeast region of the country by the third author. ...
Between October 2009 and December 2011, five Malaise traps/area were used to survey of the lance lacewings (Neuroptera: Osmylidae) in five areas of the Atlantic rainforest in São Paulo state, Brazil. Were obtained 73 specimens of Isostenosmylus pulverulentus (Gerstaecker, 1894) (96.1% of the total collected), two of Gumilla adspersus Navás, 1912 (2.6%), and one of Isostenosmylus sp. (1.3%). The osmylids were more frequently observed in the Parque Estadual da Serra do Mar / Núcleo Santa Virgínia (59.2% of the total collected), Parque Estadual Intervales (30.3%) and Parque Estadual Morro do Diabo (7.9%), inland collection sites in the state of São Paulo, Brazil. The geographic range of G. adspersus is expanded to include the states of São Paulo and Rio de Janeiro, Brazil, based on two studied specimens and one record in the citizen science website iNaturalist.
... The terminology of wing venations generally follows Ardila-Camacho et al. (2021). Although some authors interpreted the posterior-most branch of the RP (radius posterior) as MA (media anterior) in both fore-and hindwing (e.g., Aspöck et al. 1980;Lambkin 1986), we here consider it as a part of the RP, following Breitkreuz et al. (2017) and Winterton et al. (2019). The abbreviations of morphological terms used for wing venation are as follows: A, anal vein; A1-A3, first to third anal vein; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; M, media; MA, media anterior; MP, media posterior; R, radius; RA, radius anterior; RP, radius posterior; rt, radial triangle; Sc, subcostal posterior. ...
Japanese Mantispidae are revised, redescribed and illustrated with details of male and female terminalia. The following seven species are recognised: Austroclimaciella quadrituberculata (Westwood, 1852); Euclimacia badia Okamoto, 1910; Eumantispa harmandi (Navás, 1909); Tuberonotha strenua (Gerstaecker, 1894); Necyla shirozui (Nakahara, 1961); Mantispilla japonica (McLachlan, 1875); and M. transversa Stitz, 1913. In addition, we propose synonymising A. habutsuella (Okamoto, 1910) and A. subfusca (Nakahara, 1912) with A. quadrituberculata and M. j. diminuta Matsumura, 1907 with M. japonica.
... Lance lacewings (Neuroptera: Osmylidae) are an archaic group, morphologically retaining some plesiomorphic characters that are absent in most neuropterans, such as the three ocelli (in adults of most species), the nygmata, and the female gonostyli 9 (Winterton et al., 2019). Considering the mitochondrial genomic structure, Osmylidae retains the ancestral arrangement of the mitochondrial genes among the few lacewing families . ...
... Osmylidae has a diverse fossil record extending to the Lower Jurassic (Winterton et al., 2019), with 64 species in 38 genera (Makarkin et al., 2021;Ma et al., 2022). However, compared with the rich compression fossils of Osmylidae, the records of lance lacewings in amber are extremely rare. ...
... However, compared with the rich compression fossils of Osmylidae, the records of lance lacewings in amber are extremely rare. Currently, there are only two species of Osmylidae known from the amber inclusion, i.e., Nuddsia magnifica (Myskowiak et al., 2016) (subfamily Gumillinae) from the mid-Cretaceous Myanmar amber (Myskowiak et al., 2016), and Osmylidia picta (Hagen in Pictet-Baraban and Hagen, 1856) (subfamily Protosmylinae) from the Eocene Baltic amber (Winterton et al., 2019;Lu and Liu, 2021;Makarkin et al., 2021). Nevertheless, a notably rich osmylid-like larvae had been found from the Cretaceous Myanmar amber (Haug et al., 2021). ...
The family Osmylidae is an archaic group of Neuroptera and has rich fossil records. Here we present a review of Osmylidae from the mid-Cretaceous amber of northern Myanmar. We describe a new species: Nuddsia simplex sp. nov., and a new genus with one new species: Myanmarosmylus wintertoni gen. et sp. nov. Supplementary description of Nuddsia magnifica (Myskowiak et al., 2016) is also provided, with the male genitalia described for the first time. Myanmarosmylus wintertoni gen. et sp. nov. represents the first record of the subfamily Protosmylinae from the Cretaceous amber. Our new findings provide important evidence for understanding the palaeodiversity and morphology of the Cretaceous lance lacewings.
... Drawings were vectorized from photographs using the software Adobe Illustrator CS®. Morphological terminology follows Martins et al. (2019) and Winterton et al. (2019), genital sclerites were colored with the same color used by these authors, i.e. to females: yellow to sternite 8, green to gonapophyses 9, blue to tergite 9 and orange to gonocoxite 9. Wing venation terminology was based on Breitkreuz et al. (2017). A distribution map was produced using the website https://www.simplemappr.net/. ...
... Generic characters. See Martins et al. (2019) and Winterton et al. (2019). Martins et al. 2019, I. julianae Aspöck et al. 2016, I. morenoi (Navás, 1928, I. nigrifrons Kimmins, 1940, I. penai Machado et al. 2019, I. pulverulentus (Gerstaecker, 1894 Diagnosis. ...
The genus Isostenosmylus Krüger, 1913 contains 18 species distributed in South America, especially in the Andean region and in southern Brazil and Paraguay. Herein, a new species – Isostenosmylus derpi sp. n. – from a specific habitat named “Brejo de Altitude” from Paraíba state, Northeast Brazil, is described and illustrated. Distribution map and an updated taxonomic key for the genus females are provided.
Keywords:
Brejo de Altitude; Atlantic Forest; Caatinga province: Lance lacewing; Stenosmylinae
... The diversity, abundance and distribution of Neuroptera in forests can be affected by habitat types and forms, and the altitude of these environments can influence the fauna of some families (Bozdoğan 2020;Cancino-López et al. 2022). Osmylids can be found in humid habitats close to aquatic ecosystems (e.g., Osmylinae and Kemypninae) or living in leaf litter and under bark (e.g., Eidoporisminae, Porisminae, and Stenosmylinae) (New 1974;Cover and Resh 2008;Winterton et al. 2010Winterton et al. , 2019. ...
... Osmylidae (Neuroptera), commonly known as lance lacewings, are present in almost all biogeographical regions, except for the Nearctic, where fossil records indicate that the family has recently gone extinct there (Oswald 1994;Shepard and Contreras-Ramos 2009;Ardila-Camacho and Noriega 2014;Martins et al. 2016;Winterton and Wang 2016;Winterton et al. 2019). According to Shepard and Contreras-Ramos (2009), Osmylidae and other aquatic Neuroptera have no economic importance, not even as bioindicators, nevertheless, this still needs to be confirmed with further studies. ...
... There are about 280 described species of Osmylidae distributed in 25 living and 38 extinct genera (Winterton et al. 2019 Navás, 1912 (Gumillinae) are known from Brazil Martins et al. 2016Martins et al. , 2019Machado and Martins 2022), all of them from the Atlantic rainforest biome in the South and Southeast regions of the country (Machado and Martins 2022). As large territorial extensions of the country remain little explored, the fauna of Osmylidae in many Brazilian biomes is certainly underrepresented, for example, a new species of Isostenosmylus is being described to northeast region of the country by the third author. ...
Between October 2009 and December 2011, five Malaise traps/area were used to survey of the lance lacewings (Neuroptera: Osmylidae) in five areas of the Atlantic rainforest in São Paulo state, Brazil. Were obtained 73 specimens of Isostenosmylus pulverulentus (Gerstaecker, 1894) (96.1% of the total collected), two of Gumilla adspersus Navás, 1912 (2.6%), and one of Isostenosmylus sp. (1.3%). The osmylids were more frequently observed in the Parque Estadual da Serra do Mar / Núcleo Santa Virgínia (59.2% of the total collected), Parque Estadual Intervales (30.3%) and Parque Estadual Morro do Diabo (7.9%), inland collection sites in the state of São Paulo, Brazil. The geographic range of G. adspersus is expanded to include the states of São Paulo and Rio de Janeiro, Brazil, based on two studied specimens and one record in the citizen science website iNaturalist.
... Despite the low species diversity of the family in the country, the Brazilian taxa are very important to the phylogenetic studies of the family evolution, especially due to the presence of Gumilla, the sole extant genus classified in the subfamily Gumillinae, and known to possess many plesiomorphic characters . The phylogeny of the family was studied recently by Winterton et al. (2017) and a list of all extant and extinct species was presented by Winterton et al. (2019a). The Brazilian species can be identified by the papers by . ...
... Araripeberotha fairchildi Martins-Neto & Vulcano, 1990a Caririberotha fairchildi (Martins-Neto & Vulcano, 1990a Family Cratosmylidae Myskowiak, Escuillié & Nel, 2015 Taxonomic notes. The taxonomic status of Cratosmylidae is unsettled in recent literature, with some authors recognizing Cratosmylidae as a separate family (e.g., Makarkin et al., 2017;Archibald and Makarkin, 2020;, and other authors incorporating 'cratosmylid' taxa into different circumscriptions of Nymphidae (e.g., Menon et al., 2005;Winterton et al., 2019). While the matter is currently not well resolved, herein, we treat Araripenymphes and Cratosmylus in Cratosmylidae following . ...
The currently-known extinct Neuropterida fauna of Brazil comprises 18 families, 55 genera and 99 species (Neuroptera: 15 families, 49 genera, 89 species; Raphidioptera: 1 family, 2 genera, 5 species, + 1 genus incertae sedis with 2 species; Megaloptera: 1 family, 2 genera, 2 species; Order incertae sedis: 1 family, 1 genus, 1 species). All of these species have been described since 1980 and they collectively constitute the overwhelming majority of extinct Neuropterida presently known from South America. Other than one species of Permithonidae (Neuropterida incertae sedis) described from the Late Permian Irati Formation (Paraná Basin) of Rio Grande do Sul state, all extinct Brazilian Neuropterida taxa are known only from the Early Cretaceous Crato Formation (Araripe Basin) in Ceará and Pernambuco states. Of the 35 authors who have contributed to the descriptive literature of this fauna over the past 42 years, the work of Brazilian paleontologists Rafael Gioia Martins-Neto [1954-2010] and Maria Aparecida Vulcano-d'Andretta [1921-2018] - who together contributed to the description of 67 species - is especially notable. However, since their deaths, extinct Neuropterida have received little attention from Brazilian scientists and most later work on this fauna has been undertaken by workers outside Brazil. We provide this updated synthesis of the extinct Neuropterida of Brazil in order to centralize baseline knowledge for this fauna, to support efforts to document and conserve Brazilian fossil sites, and to encourage future work on the fauna, particularly by the next generation of Brazilian paleoentomologists.
Keywords:
Fossil; Crato formation; Irati formation; Taxonomy
... Fossil neuropterans have been found in quite large numbers for adults (e.g. Engel & Grimaldi 2007;Menon & Makarkin 2008;Jepson 2015;Winterton et al. 2019), but also for larvae (e.g. Pérez-de la Fuente et al. 2020;Haug et al. 2021a), while fossil pupae are an absolute rarity (see discussion). ...
During ontogenetic development, insects can undergo quite drastic changes (metamorphosis) until the adult stage is reached. A substantial part of this development in one group of Insecta, Holometabola, takes place during the pupa stage. Despite the pupa being recognised as an important phase, rather few depictions of pupae exist in the literature. We report here the first find of a fossil pupa of the lacewing group Mantispidae. The specimen represents an exuvia and is enclosed in Ukrainian Rovno amber, Eocene in age (c. 35–40 million years). We review the entire record of extant pupae of Mantispidae depicted in the literature or in online image repositories. With the aid of elliptic Fourier analysis, we compare the outline of the femur of the foreleg (raptorial appendage in the adults) of pupae and adults of Mantispidae. The pupae are all very similar concerning the femur, while the adults show a larger morphological diversity, particularly the extinct forms. Furthermore, our results indicate that the forelegs do not become increasingly complex throughout ontogenetic stages, but instead undergo an indirect development. According to the low variation in morphology seen in the pupa stage in Mantispidae, it is plausible that it represents a phylotypic stage for the group, i.e. a phase characterised by a significantly lower variability than other stages.
... nov., from the Middle Jurassic Jiulongshan Formation in Daohugou, Inner Mongolia, China. This new species can be assigned to Kempyninae based on the following apomorphic characters: The M space in the hind wing is extremely broad and sinuate cross-veins are arranged in the intramedial area to produce two rows of irregular cells, which are regarded as an apomorphy of Kempyninae (Winterton et al., 2019;Ma et al., 2020a). To investigate the inner relationships of Kempyninae, we conducted a phylogenetic analysis including all extant and most fossil genera. ...
... The line drawing was produced using the Adobe Photoshop CC and Adobe Illustrator CC software. The terminology for wing venation and genitalia follows Breitkreuz et al. (2017) and Winterton et al. (2019). ...
... Remarks. The hind wing is significant to the generic taxonomy of Kempyninae, and many fossil genera were established based on the characters of the hind wing (Winterton et al., 2019). The genus Arbusella was first erected by Khramov based on a species from the Late Jurassic of Kazakhstan, which was most characterized by the presence of a row of gradate series inbetween subcostal veinlets (Khramov, 2014a). ...
The extant kempynines, a strict “southern group,” are confined to South America and Australia, while their most fossil relatives are abundantly recorded in the Northern Hemisphere. This pattern of the biogeographic distribution implies the complicated evolutionary scenario of Kempyninae. Herein, a new northern species Arbusella platyptera Ma et Wang, sp. nov. is described from the Jiulongshan Formation in Daohugou, Inner Mongolia, China. Additionally, a key to the extinct species and extant genera of Kempyninae is provided. Integrating all extant and most fossil genera of Kempyninae, we conducted phylogenetic analyses to explore the inner relationships of Kempyninae for the first time. The results corroborate the monophyly of Kempyninae and retrieve three clades within the subfamily, namely, two northern fossil genera (†Arbusellla + †Jurakempynus), constituting the basalmost clade and three other northern fossil genera (†Sauktangida + †Mirokempynus + †Ponomarenkius), forming a monophylic clade, which is sister to the third clade that includes all extant southern genera and the southern fossil genus of †Euporismites. Also, the extant kempynines were hypothesized to evolve independently from their northern Mesozoic relatives. The Dispersal-vicariance (DIVA) analysis revealed a northern and prepangean origin of Kempyninae, and the northern ancestral kempynines first colonized the Southern Hemisphere before the split of Pangea. Our results expose a more complicated evolutionary scenario of the insects with a long evolutionary history and provide new insights into the formation of distribution patterns in current relictual insects.
... Cratosmylidae is among the extinct myrmeleontoid familes with questionable familial status, possibly being paraphyletic or a stem group in Myrmeleontoidae [23,35]. The two cratosmylid genera Araripenymphes and Cratosmylus were once placed within the family Nymphidae, and the latter genus was originally placed within the family Osmylidae [13,26,36]. The close relationship of Cratosmylidae and Babinskaiidae was first proposed by Makarkin et al. [23], but no argument was given. ...
The extinct neuropteran families Cratosmylidae and Babinskaiidae hitherto only known from the Cretaceous represent the transitional lineage between Nymphidae and advanced myrmeleontoids (e.g., Nemopteridae and Myrmeleontidae) in the superfamily Myrmeleontoidea. Here, we describe two new species, which respectively belong to Cratosmylidae and Babinskaiidae, namely, Araripenymphes burmanus sp. nov. and Paradoxoleon chenruii gen. et sp. nov., from the mid-Cretaceous Kachin amber of Myanmar. Cratosmylidae, which was previously only recorded from the Lower Cretaceous of Brazil (Crato Formation), is first reported from the mid-Cretaceous Kachin amber of Myanmar, and the co-occurrence of Araripenymphes Menon, Martins-Neto and Martill, 2005 across South America and Asia further documents the Gondwanan origin of the northern Myanmar amber lacewing paleofauna. The first finding of a deeply bifurcated forewing MP with two free branches in Babinskaiidae (viz., Paradoxoleon chenruii gen. et sp. nov.) highlights the morphological diversity of this extinct family. The phylogenetic positions of Araripenymphes burmanus sp. nov. and Paradoxoleon chenruii gen. et sp. nov. were recovered on the basis of a morphology-based phylogenetic analysis, and the monophyly of Cratosmylidae + Babinskaiidae was corroborated. Given the paraphyly of Cratosmylidae, its familial status is discussed.
