A camouflaged diversity: taxonomic revision of the thorny lacewing subfamily Symphrasinae (Neuroptera, Rhachiberothidae).

Species of the thorny lacewing subfamily Symphrasinae (Neuroptera: Rhachiberothidae) are revised. Prior to this work, 42 species were known in the genera Anchieta Navás, 1909, Plega Navás, 1928, and Trichoscelia Westwood, 1852. Herein, the number of species is increased to 60, 23 of which are newly described. Species previously known are redescribed, and their taxonomic status is revised. Keys, diagnoses, and high-resolution images for all species are presented. The distribution range of Anchieta is now known from Costa Rica to southern Brazil with a total of 11 species, of which three are newly described. The genus Plega is known from southwestern United States to southern Brazil and includes 28 species of which 14 are described as new. Moreover, the genus Trichoscelia occurs from central and southern Mexico to Argentina, with a total of 21 species, of which six are herein newly described. A phylogenetic analysis of Symphrasinae based on morphological characters recovered the three symphrasine genera as monophyletic, with Anchieta sister to Plega + Trichoscelia. The three genera are newly diagnosed based on a cladistic framework. Within the genus Anchieta, bee-mimicking species comprise a monophyletic group, while wasp-mimicking species form a laddered sequence to that lineage. Within Plega, three lineages are recovered, the first mostly composed of South and Mesoamerican species, the second with species predominantly from Central America and central and southern Mexico, and a third clade encompassing species mostly from central and northern Mexico and southwestern United States. By contrast, relationships between species of Trichoscelia were poorly resolved because of a simplified and conserved morphology of this group.

Comparative morphology of male genitalia in antlions (Insecta, Neuroptera, Myrmeleontidae), with emphasis on owlflies (Ascalaphinae) and a possible structural evolutionary scenario.

Male genitalia morphology in Myrmeleontidae has traditionally been insufficiently studied, although it has received increased attention for its diagnostic value in recent times. A neutral terminology has generally been applied in standard taxonomic practice, yet knowledge of an equivalent and stable terminology across taxa based on comparative morphology has been missing. Herein a detailed comparative morphology study with examples from most tribes within Myrmeleontidae, including owlflies (Ascalaphinae), attempts to relate external and internal genital structures based on a proposed groundplan for Neuroptera and Myrmeleontidae. We contend that a groundplan based on 10 abdominal segments, plus vestigial structures from an 11th segment, coherently depicts structural components across myrmeleontid taxa. A gonarcus, an element of Neuropterida amply referred in Neuroptera, is supported to represent the pair of abdominal appendages of segment X medially fused, with gonocoxite and gonostylus components. In most myrmeleontid taxa, basal (gonocoxites) and distal (gonostyli) components separate, with gonostyli positioned posteriorly with respect to gonocoxites, still united with translucent, lightly sclerotized tissue, forming a more or less conical structure, a proposed synapomorphy for the family. Ninth gonostyli are generally reduced (pulvini) and have migrated close to the base of gonarcus (10th gonocoxites). A pelta, also a potential synapomorphy for Myrmeleontidae, derives from paired setose surfaces of the 10th gonostyli, medially positioned (still evident in Bubopsis). Three structural types of gonarcus are diagnosed for illustrative purposes, as they may represent convergent constructs.

The genus Climaciella Enderlein, 1910 (Neuroptera, Mantispidae) in French Guiana.

The genus Climaciella Enderlein, 1910 is a remarkable group of mantidflies (Neuroptera: Mantispidae: Mantispinae) distributed from Canada to Argentina, including parts of the Caribbean. This genus comprises nine valid extant species plus an extinct species from the late Oligocene of France. Species exhibit Batesian mimicry with vespid wasps (Vespidae). Herein, six species of Climaciella from French Guiana are documented. Before this study only C.semihyalina (Le Peletier de Saint Fargeau & Audinet-Serville in Latreille et al. 1825) was known from this territory. Two new species, C.elektroptera Ardila-Camacho, Winterton & Contreras-Ramos, sp. nov. and C.nigriflava Ardila-Camacho, Winterton & Contreras-Ramos, sp. nov., are described as well as the first records of C.amapaensis Penny, 1982, and C.tincta (Navás, 1914) provided from French Guiana. An unknown species recorded from a single female specimen is also presented. Based on the examination of material of C.amapaensis recorded here, a specimen previously recorded from Colombia as belonging to this species is herein proposed as a new species, C.risaraldensis Ardila-Camacho, sp. nov. A taxonomic key and high-resolution images of the species from French Guiana are provided.

Antlions of formerly recognized tribe Gnopholeontini (Neuroptera: Myrmeleontidae: Brachynemurini) from Peninsula of Baja California, with a new species of Tyttholeon Adams

ABSTRACT A new antlion species, Tyttholeon froehlichi Tavares, Marquez and Contreras sp. n., is described from the Baja California Peninsula, Mexico, an area biogeographically significant for its high number of endemic species. Previously monotypic, Tyttholeon Adams now comprises two Nearctic species. Three of the four genera within the formerly recognized tribe Gnopholeontini and their respective species, all present in the Peninsula of Baja California (Gnopholeon barberi Currie, G. delicatulus Currie, G. zapotecus Stange, Menkeleon bellulus Banks, and Tyttholeon puerilis Adams), are herein diagnosed, illustrated, and have their distribution updated and analyzed. A taxonomic key is provided for Tyttholeon.

Diversity of Lacewings (Neuroptera) in an Altitudinal Gradient of the Tacaná Volcano, Southern Mexico.

Neuroptera is an order of insects with a moderate diversity of species numbers yet a high between-family morphological diversity, which has a significant ecological role as a predator. However, there are few studies focused on describing changes in species diversity along environmental gradients. We evaluated changes in the alpha and beta diversity of species and the higher taxa in Neuroptera communities in the Tacaná Volcano in southern Mexico. Five sites each at different altitudes were studied through systematic annual sampling. The taxonomic and phylogenetic alpha diversity were analyzed, as well as the beta diversity and its components, species turnover and nestedness. The alpha diversity had two trends: (1) decreased standardized richness and taxonomic distinctness with increasing altitude, and (2) increased estimated richness and species diversity at intermediate altitudes. The highest turnover values for species, as well as for supra-specific taxa, were recorded at sites with lower altitudes. The highest total beta diversity value was recorded at elevations above 3000 m, whereas the highest number of species and supra-specific taxa were observed at sites between 600 and 2000 m, with an evident decrease above 3000 m. The type of vegetation and environmental conditions may be influencing the decrease in diversity toward higher elevations, which could explain the niche specialization of Neuroptera species to particular sites within the gradient. These results highlight the need to study the environmental factors and their effects on species composition along an elevation gradient.

Similar pattern, different paths: tracing the biogeographical history of Megaloptera (Insecta: Neuropterida) using mitochondrial phylogenomics.

The sequential breakup of the supercontinent Pangaea since the Middle Jurassic is one of the crucial factors that has driven the biogeographical patterns of terrestrial biotas. Despite decades of effort searching for concordant patterns between diversification and continental fragmentation among taxonomic groups, increasing evidence has revealed more complex and idiosyncratic scenarios resulting from a mixture of vicariance, dispersal and extinction. Aquatic insects with discreet ecological requirements, low vagility and disjunct distributions represent a valuable model for testing biogeographical hypotheses by reconstructing their distribution patterns and temporal divergences. Insects of the order Megaloptera have exclusively aquatic larvae, their adults have low vagility, and the group has a highly disjunct geographical distribution. Here we present a comprehensive phylogeny of Megaloptera based on a large-scale mitochondrial genome sequencing of 99 species representing >90% of the world genera from all major biogeographical regions. Molecular dating suggests that the deep divergence within Megaloptera pre-dates the breakup of Pangaea. Subsequently, the intergeneric divergences within Corydalinae (dobsonflies), Chauliodinae (fishflies) and Sialidae (alderflies) might have been driven by both vicariance and dispersal correlated with the shifting continent during the Cretaceous, but with strikingly different and incongruent biogeographical signals. The austral distribution of many corydalids appears to be a result of colonization from Eurasia through southward dispersal across Europe and Africa during the Cretaceous, whereas a nearly contemporaneous dispersal via northward rafting of Gondwanan landmasses may account for the colonization of extant Eurasian alderflies from the south.

Aquatic beetle diversity from Volcán Tacaná, Mexico: altitudinal distribution pattern and biogeographical affinity of the fauna.

Results of an aquatic beetle survey at Volcán Tacaná, Mexico, are presented with five altitudinal levels in a monthly sampling regime, aiming to estimate both diversity and altitudinal distribution patterns of the aquatic beetle fauna. The first list of aquatic beetle species from this mountain is presented, comprising 40 species in 32 genera and nine families, with four species recorded for the first time from Mexico and six recorded for the first time from Chiapas. The aquatic beetle fauna is characterized by Elmidae with 20 species, Dytiscidae with eleven species, Dryopidae with three, and Epimetopidae, Hydraenidae, Hydrophilidae, Gyrinidae, Lutrochidae, and Noteridae with one species each. The species composition through the sampled altitudinal gradient (670-1,776 m) was not homogeneous, with the elmid genera Macrelmis, Heterelmis, Microcylloepus, and Austrolimnius present at all levels, while Hexanchorus, Neoelmis, and Onychelmis were present at levels 1-3 (673-1,214 m); dytiscids were mostly present at levels 4 and 5 (1,619-1,776 m), and dryopids were present only at levels 1-3. A Parsimony Analysis of Endemicity supports a general partition between altitudinal levels 1-3 and levels 4 + 5.

Larval and female descriptions of Mejicanotrichia Harris & Holzenthal, 1997 (Trichoptera, Hydroptilidae, Leucotrichiinae) from Mexico.

Mejicanotrichia Harris & Holzenthal, 1997 is a small genus of Hydroptilidae (Trichoptera), which consists of seven species, six of them distributed in Mexico, and one more in Guatemala. Larval descriptions of only two species (M.blantoni and M.estaquillosa) were previously known, as well as only females of three species (M.blantoni, M.estaquillosa, and M.tamaza) previously described. The present study provides descriptions of the larvae of M.harrisi and M.tridentata, as well as a description of the female of M.harrisi. Identification keys for adult males, known females, and known larvae are also provided. This work aims to incorporate more information into the taxonomy of the genus, its ecology, and facilitate additional characters of potential use in future phylogenetic studies.

After a decade, a new Venezuelan species of Corydalus Latreille (Megaloptera, Corydalidae, Corydalinae) is discovered.

A new species of dobsonfly from Venezuela, Corydalusralphi Martins, Azevêdo, Hamada & Contreras, sp. nov., was discovered a decade after the last description of a species of this genus in the country. The new species is morphologically similar to C.wanningeri Contreras-Ramos & von der Dunk, sharing a uniform reddish coloration of body and wings and similar male genitalic structures. Likewise, it shares this particular coloration with C.neblinensis Contreras-Ramos but the genitalic structure fits within the C.crossi Contreras-Ramos species group. Two specimens, one male and one female, were collected on Tarotá River, in the Gran Sabana region, Canaima National Park, in southern Venezuela. A key to identify males of the Venezuelan species of Corydalus is provided.

First record of the genus Lainius Navás, 1913 (Neuroptera: Chrysopidae) in Mexico, with notes on the distribution of Apochrysinae

ABSTRACT The species Lainius constellatus Navás is recorded for the first time from Mexico; thus this report represents the northernmost record of the genus and species for the Americas. In addition, it provides data on the variation in color marks and a description of the hypandrium internum. A brief summary of the distribution of the species belonging to the subfamily Apochrysinae is provided, with emphasis on the three American genera, which share distribution mostly in the Pacific domain of the Brazilian subregion, ranging from the Mexican transition zone to possibly the Chacoan subregion.

A new species of Paranthaclisis Banks (Neuroptera, Myrmeleontidae) from the Peninsula of Baja California, Mexico

ABSTRACT Paranthaclisis stangei Marquez, Martins, and Contreras, sp. n., is a new myrmeleontid from Baja California Sur state, Baja California Peninsula, Mexico, which is an important area of endemism. After this new discovery, the genus Paranthaclisis is composed by five species, three occurring in Mexico. This new species is easily identified by a rostrum completely yellowish-white, an area of pre-origin of RP with spurial vein on hindwing and with marks; males are easily separated by the conspicuous posterior thinning of the parameres and mediuncus dorsally without teeth and a basomedial split.

Comparative morphology of extant raptorial Mantispoidea (Neuroptera: Mantispidae, Rhachiberothidae) suggests a non-monophyletic Mantispidae and a single origin of the raptorial condition within the superfamily.

Adult external morphology of the extant raptorial Mantispoidea (Insecta: Neuroptera: Mantispidae and Rhachiberothidae) is compared emphasizing the morphology of the subfamily Symphrasinae as a key group to understand the phylogenetic relationships among the members of the superfamily. Plega dactylota Rehn, 1939 is thoroughly characterized in order to exemplify the morphology of the Symphrasinae. Additionally, following a review of the literature and examination of comparative material of Dilaridae, Berothidae, Rhachiberothidae and all Mantispidae subfamilies, a new interpretation of the components of the raptorial apparatus (i.e., head, prothorax, grasping forelegs, as well as integumentary specializations) is presented. Also, wing venation for these groups is reinterpreted, and new homology hypotheses for wing venation are proposed based on tracheation and comparative analyses. Given the high morphological divergence on the genital sclerites within the Mantispoidea, plus the confusing previous usage of neutral terminology and terms referring to appendages across taxonomic and morphological studies, we attempt to standardize, simplify, and situate terminology in an evolutionary context under the "gonocoxite concept" (multi-coxopod hypothesis). The remarkable morphological similarity of the genital sclerites of Symphrasinae and Rhachiberothidae (sensu U. Aspöck Mansell 1994) with the Nallachinae (Dilaridae) was taken as a starting point to understand the morphology of other Mantispidae subfamilies. Based on these morphological comparisons, we provide a revised phylogenetic analysis of Mantispoidea. This new phylogenetic analysis supports a sister group relationship between the family Rhachiberothidae, comprising Rhachiberothinae and Symphrasinae, and the family Mantispidae, including the subfamily Mantispinae and its sister taxa Drepanicinae and Calomantispinae, which may represent a single subfamily. Based on these analyses, raptorial condition probably evolved a single time in these insects and subsequently became diversified in the two sister clades of the raptorial Mantispoidea.

Redescription of larva of the South African endemic dobsonfly genus Chloroniella Esben-Petersen, 1924 (Megaloptera, Corydalidae), with a proposed terminology for Corydalidae larval sclerites.

Chloroniella Esben-Petersen is a relict, monotypic genus of Corydalidae (Megaloptera), considered sister to the remaining dobsonfly genera (Corydalinae); it is the only genus of Corydalinae known from the Afrotropical region. The larva of its only species, C. peryngueyi Esben-Petersen, 1924, was originally described by Barnard in 1931. Herein we redescribe the larva of C. peryngueyi using Scanning Electron Microscopy (SEM), colored focus-stacked images, and drawings in order to provide greater morphological detail of this significant species. Also, we propose a standardized terminology for Corydalidae larval sclerites.

Comparative study of sensilla and other tegumentary structures of Myrmeleontidae larvae (Insecta, Neuroptera).

Antlion larvae have a complex tegumentary sensorial equipment. The sensilla and other kinds of larval tegumentary structures have been studied in 29 species of 18 genera within family Myrmeleontidae, all of them with certain degree of psammophilous lifestyle. The adaptations for such lifestyle are probably related to the evolutionary success of this lineage within Neuroptera. We identified eight types of sensory structures, six types of sensilla (excluding typical long bristles) and two other specialized tegumentary structures. Both sensilla and other types of structures that have been observed using scanning electron microscopy show similar patterns in terms of occurrence and density in all the studied species (with few exceptions). The sensilla identified are: coeloconica, placoidea, basiconica, trichodea type I, trichodea type II, and campaniformia. All these sensilla have mechano- or chemosensorial functions. Some regions of the larval body have been studied using SEM for the first time, such as the surface of the food canal, which bears sensilla coeloconica, and the abdominal segment X, that bears three types of sensilla: coeloconica, basiconica, and campaniformia. Sensilla placodea are newly reported on antlion larvae, being present on the mandibular base, pronotum, mentum, and cardum. Also, new locations of sensilla coeloconica (e.g., on rastra) and sensilla campaniformia (e.g., on odontoid processes) are noted. A novel porous texture with chemoreceptor function has been identified in the base of mandibles. A mechanism of dentate-notched surfaces that anchor maxillae and mandible, reinforcing the food canal, is detailed. All these sensorial structures, in addition to ocular tubercles for light caption and their great muscular system, confer to these larvae an extraordinary predation capacity to success hunting and living in such harsh environments.

A new species of Ceraeochrysa Adams (Neuroptera, Chrysopidae), with a key to the species from Mexico.

The genus Ceraeochrysa Adams is widely distributed in the New World, from southeastern Canada to Argentina, with 15 out of 61 previously known species recorded in Mexico. In this paper, Ceraeochrysa tacanensis sp. nov. is described and illustrated from Volcán Tacaná, Chiapas, and an identification key to Ceraeochrysa species present in Mexico is provided. The new species is similar to others with swollen and darkened posterior branches of the cubital vein, and it can be separated from these other species by an elongate gonapsis extending from the base of the gonosaccus; the gonapsis is slightly upturned, terminating in a rounded apex with dorsal microteeth. Females of the new species have non-distinctive genitalia morphology. However, they can be associated with males of the species by body color pattern, synchrony, and sympatry.

Two new species and a key to the dustywings of Mexico (Neuroptera, Coniopterygidae).

Two new species of Coniopterygidae are described from Mexico: Neoconis szirakii sp. n. and Coniopteryx (Scotoconiopteryx) josephus sp. n., increasing the number of recorded species of dustywings from Mexico to 43. Descriptions and illustrations are based primarily on male genitalia. A key to the known species of Coniopterygidae from Mexico is included.

Four new species of Plega Navás, 1928 (Neuroptera: Mantispidae) from Mexico.

Four new species of Plega Navás, 1928 (Mantispidae: Symphrasinae) from Mexico are herein described and illustrated, namely P. mixteca sp. n. from Oaxaca and Jalisco, P. stangei sp. n. from Morelos and Oaxaca, P. sonorae sp. n. from Sonora, and P. spinosa sp. n. from Guerrero, Jalisco, Morelos, Nayarit, Oaxaca, Puebla, and Veracruz. Including these new species, Plega's diversity raises to 17 described species, of which 12 are recorded from Mexico, and one of them, Plega variegata Navás, 1928, is herein proposed as nomen dubium. A taxonomic key to the known species of Plega from Mexico is provided.

Taxonomic review of the mantidfly genus Nolima Navás (Neuroptera, Mantispidae, Calomantispinae).

The mantidfly genus Nolima Navás, 1914 (Neuroptera, Mantispidae, Calomantispinae) is herein revised. Nolima is endemic to the New World, ranging from the southwestern United States south to Costa Rica. Nolimainfensa Navás, N.pinal Rehn, and N.victor Navás are redescribed, while the new species Nolimacostaricensis Reynoso & Contreras, sp. nov. is described from Costa Rica. The species N.dine Rehn and N.kantsi Rehn are synonymized with N.pinal. Additionally, the species N.praeliator Navás and N.pugnax Navás are synonymized with N.victor, for which a lectotype is designated. New distribution records are provided from Guatemala and Honduras for Nolimainfensa, the state of Nevada in western United States for N.pinal, and the state of Puebla in central Mexico for N.victor. An illustrated key and a distribution map are presented.

A new species of Wesmaelius Krüger from Mexico, with a key to the New World species of the subgenus Kimminsia Killington (Neuroptera, Hemerobiidae).

Wesmaelius (Kimminsia) nanacamilpa Marquez & Contreras, sp. n., a brown lacewing from Tlaxcala state, Mexico is described and illustrated. This is the second recorded species of Wesmaelius from Mexico, and the third from Middle America. Males of the new species may be identified by parameres separate apically, styliform sclerites directed basally, as well as a rounded gonarcus with a short entoprocessus. Females may be distinguished from closely related species by a subgenital plate with the central plate broadly incised basally. There are now 16 species of Wesmaelius known from the New World.

Bionomics and Ecological Services of Megaloptera Larvae (Dobsonflies, Fishflies, Alderflies).

Megaloptera belong to a large monophyletic group, the Neuropteroidea, together with Coleoptera, Strepsiptera, Raphidioptera, and Neuroptera. With the latter two, this order constitutes the Neuropterida, a smaller monophyletic subset among which it is the only entirely aquatic group, with larvae of all species requiring submersion in freshwater. Megaloptera is arguably the oldest extant clade of Holometabola with aquatic representatives, having originated during the Permian before the fragmentation of Pangea, since about 230 Ma. It includes 54 genera (35 extant and 19 extinct genera), with 397 extant described species and subspecies. Recent Megaloptera are divided into two families: Corydalidae (with subfamilies Corydalinae-dobsonflies and Chauliodinae-fishflies) and Sialidae (alderflies), both widely yet disjunctively distributed among zoogeographical realms. All species of Megaloptera have aquatic larvae, whereas eggs, pupae, and adults are terrestrial. The anatomy, physiology, and behavior of megalopteran larvae are specialized for an aquatic predatory habit, yet their ecological significance might still be underappreciated, as their role in food webs of benthic communities of many temperate and tropical streams and rivers is still understudied and largely unquantified. In many freshwater ecosystems, Megaloptera larvae are a conspicuous benthic component, important in energy flow, recycling of materials, and food web dynamics.