Third instar of the myrmecophilous Italochrysa insignis (Walker) from Australia (Neuroptera: Chrysopidae: Belonopterygini).

The third instar of the Australian green lacewing Italochrysa insignis (Walker) is described and compared with that of the European Italochrysa italica (Rossi). Larvae of both species are associated with arboreal ant nests. Moreover, they share a substantial number of morphological adaptations that may defend them against ants and that distinguish them from larvae of other chrysopids. The larvae of the two Italochrysa Principi species also have distinct differences. The potential systematic value of several of the larval characters at the tribal, generic, and species levels is assessed, and a brief review of myrmecophily in the Belonopterygini is presented.

Adults and larvae of two Leucochrysa (Leucochrysa) species (Neuroptera: Chrysopidae): descriptions, biological notes, and relationships.

This taxonomic study includes: (i) a redescription of Leucochrysa (Leucochrysa) nigrilabris (Banks) (♂ and ♀), based on the type specimen and new material and (ii) images of the Leucochrysa (L.) insularis (Walker) type, adult color polymorphism, and genital characters (♂ and ♀). For both species, it provides: (iii) descriptions of the larvae, (iv) biological notes, and (v) geographic records. Using the above information, we compare the two species with each other and with other Leucochrysa (Leucochrysa) species that purportedly are closely related. We conclude: First, the larval features of L. (L.) nigrilabris and L. (L.) insularis coincide with those previously described as characteristic of the genus Leucochrysa and its subgenus Leucochrysa. Second, based on their genitalia (♂ and ♀), larval morphology, and biology (e.g., deposition of eggs in clusters), the two species are distinct, but very closely related. And, third, L. (L.) nigrilabris and L. (L.) insularis share several characteristics with the Leucochrysa (L.) "varia-like" species; these include similar adult color polymorphisms and aspects of their larval morphology. However, their genitalia (♂ and ♀) differ significantly from those of the described L. (L.) "varia-like" species, and thus we consider the two sets of species to be distinct.

Discovery and redescription of the true Nuvolumbrosus Navás and naming of a new Nuvol species (Neuroptera, Chrysopidae, Leucochrysini).

Examination of a newly discovered specimen of Nuvol showed that our earlier species determination of Nuvolumbrosus Navás had been incorrect and that our "redescription" of the species actually applied to an undescribed species. Here, we redescribe the true N.umbrosus, based on a newly discovered male specimen. This specimen closely resembles Navás' description, and it was collected from the Atlantic Forest as was the original type specimen. In addition, we assign the previously misidentified Nuvol specimens from the Amazonian region to a separate species, Nuvolsatur Sosa & Tauber, sp. nov. As a result of these actions, the genus Nuvol now contains two morphologically and geographically distinct species. In addition, the abdomens and genitalia of both sexes of Nuvol are now described (although each from a separate species).

The New World Belonopterygini (Neuroptera: Chrysopidae): descriptions of a new genus and species from the West Indies and comparisons among the genera.

Pantaleonius toschiorum Tauber, n. gen., n.sp., is described from several islands in the Bahamas and Greater Antilles of the West Indies. For comparison, Abachrysa eureka Banks from southeastern United States is redescribed, and data on the three remaining New World belonopterygine genera, Vieira Navás, Nacarina Navás, and Belonopteryx Gerstaecker, are summarized. A key for identifying the New World belonopteryine genera is included. Although recognizably distinct, Pantaleonius shares several features with Vieira, which is considered basal within Belonopterygini. Both genera are quite different from Abachrysa Banks, which is known to be derived among the New World belonopterygines and also closely related to the Old World belonopterygine clade.

The Neotropical green lacewing genus Ceraeochrysa Adams (Neuroptera: Chrysopidae)-new synonymies and combinations, a new species, and an updated key to species.

Ceraeochrysa Adams, 1982, with ~60 species, is a diverse and relatively well studied New World genus of green lacewings. However, nomenclatural problems and misidentifications persist; undescribed species continue to be found; and species determinations remain difficult. Here, we address several problematic issues within the genus, and we justify the following taxonomic changes: Part 1 synonymizes Ceraeochrysa rafaeli Adams Penny syn. n. and Chrysopodes (Chrysopodes) nigropictus Freitas Penny syn. n. with Ceraeochrysa discolor (Navás). Part 2 identifies Ceraeochrysa adynatos (Freitas Penny) comb. n. as the new name for Chrysopodes (Chrysopodes) adynatos Freitas Penny. Part 3 renames Chrysopa sarta Banks as Ceraeochrysa sarta (Banks) comb. n, and identifies it as conspecific with Ceraeochrysa berlandi (Navás) syn. n. Part 4 names Chrysopa poujadei Navás as Ceraeochrysa poujadei (Navás) comb. n. and also identifies its new synonyms-Chrysopodes (Chrysopodes) laevus (Navás) syn. n. and Ceraeochrysa forcipata Freitas Penny syn. n. Part 5 redescribes the enigmatic Ceraeochrysa silvanoi (Navás) and recognizes Ceraeochrysa tucumana (Navás) syn. n. as its junior synonym. Part 6 describes a new Ceraeochrysa species: Ceraeochrysa paraensis Sosa sp. n. from Brazil. Finally, to help facilitate species identifications, Part 7 incorporates all new species and all taxonomic changes made since the last revision of the genus (2009) into an updated key to the currently valid species of Ceraeochrysa. Lectotypes are designated for three species: C. discolor Navás, C. sarta Banks, and C. tucumana Navás.

Larvae of Abachrysa eureka (Banks) (Neuroptera: Chrysopidae: Belonopterygini): descriptions and a discussion of the evolution of myrmecophily in Chrysopidae.

Here we describe the first and third instars and the egg of the New World chrysopid Abachrysa eureka (Banks). Like other members of the tribe Belonopterygini, this species is myrmecophilic. Comparisons of Abachrysa larval and egg characteristics with those reported from four other belonopterygine genera indicate that Abachrysa more closely resembles the Old World Calochrysa and Italochrysa than the New World Vieira and Nacarina. The three genera Abachrysa, Calochrysa and Italochrysa all have large eggs, accelerated embryonic development, and an elaborate set of morphological modifications for larval debris carrying, an important defense against ant attack. This pattern of shared features is consistent with the phylogenies recovered in recent molecular studies that place the New World genera Vieira and Nacarina basal to Abachrysa and the Old World genera. Our assessment of current morphological information in relation to the molecular studies indicates the following sequence: (i) The form of belonopterygine myrmecophily that is currently expressed in the basal lineages (Nacarina and perhaps Vieira) originated in the New World and does not involve elaborate larval modifications for debris carrying. (ii) Myrmecophily that is based on a correlated set of developmental and morphological traits subserving debris carrying evolved in the New World when Abachrysa diverged. (iii) Subsequently, the debris-carrying clade of Belonopterygini underwent a significant radiation in the Old World, but not in the New World.

South American Nothochrysinae (Neuroptera, Chrysopidae): I. Description of Nothochrysaehrenbergi sp. nov.

A new species, Nothochrysaehrenbergi sp. nov., is described from Chile; it is the first species of Nothochrysa to be reported from the Southern Hemisphere and only the second from the New World. The genus now contains six extant species as well as two species known from late Oligocene and Miocene fossils. An updated catalog of the valid Nothochrysa species is presented, and three nomina dubia are discussed. The inclusion of the new species in Nothochrysa is well supported by morphological features. However, it and other species currently in the genus also share significant features with Archaeochrysa, an older genus of Nothochrysinae which is known only from the Eocene (Ypresian) to the late Oligocene. It therefore appears that N.ehrenbergi is among the least derived Nothochrysa species, and that the separation of Archaeochrysa from Nothochrysa is open to question and further examination.

South American Nothochrysinae (Neuroptera, Chrysopidae): II. Redescription of Leptochrysaprisca Adams & Penny.

Leptochrysa Adams & Penny is one of four genera of Nothochrysinae recorded from the New World. Previously, this genus and its only described species, Leptochrysaprisca Adams & Penny, were known from a single female specimen that is discolored and damaged by fungal infestation. Thus, accurate information on the taxon was limited mostly to the wings and some other external features. Here, I describe a recently collected, second female specimen with the goal of providing images of the adult coloration and elucidating characters (especially the female genitalia) that were unavailable earlier. Some variation between the two known specimens is also noted and used in interpreting venation characters. Finally, comparisons are made with other extant genera pertaining to the placement of the species within Chrysopidae.

A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters.

A new and very unusual species of green lacewings, Ungla pseudomeleoma sp. nov. is described from Peru. Males of this species exhibit a remarkable example of generic-level homoplasy with Meleoma. Its abdominal and genitalic features are completely consistent with the genus Ungla. However, its external sexually dimorphic features resemble those of both Ungla and Meleoma. Females of the new species also express cranial modifications that resemble those of some female Meleoma species.

Type specimens of Neuropterida in the Hope Entomological Collection, Oxford University Museum of Natural History.

The Hope Entomological Collection in the Oxford University Museum of Natural History holds a large and diverse array of historically valuable type specimens for species in the superorder Neuropterida (Megaloptera, Neuroptera, and Raphidioptera). Most are from the mid-1800s (1823-1874). Here, we report each type found during a rigorous search of the collection, and we confirm the identity and status of each type with bibliographic, specimen, and label data. Images, current nomenclatural information, and references for name changes are provided for each species. We identified primary or secondary types for 76 species in seven families of Neuroptera, primary and secondary types for one species of Raphidoptera (Raphidiidae), and secondary types for three species of Megaloptera (Corydalidae). Among the Neuroptera, we found primary types for 26 species of Mantispidae, 16 species of Myrmeleontidae, 11 species of Ascalaphidae, seven species of Nemopteridae, five species of Chrysopidae, and one species each of Coniopterygidae and Hemerobiidae. Types for only two species that were reported to be in the collection were not found. To help stabilize the nomenclature, we made new lectotype designations for the following six species in the Hope Collection: Colobopterusdissimilis McLachlan, 1871; Mantispabatesella Westwood, 1867; Mantispamyrapetrella Westwood, 1867; Mantispatropica Westwood, 1852; Myrmeleonanomalus Rambur, 1842; and Myrmeleonsingulare Westwood, 1847. We also made new lectotype designations for two species in the Natural History Museum in London: Mantispaindica Westwood, 1852, and Mantispaquadrituberculata Westwood, 1852. In several other cases, holotype determinations in the literature were recognized as lectotype designations. Finally, to correct an important error in the literature, we reinstated the synonymization of Ascalaphusanticus Walker, 1853, A.loquax Walker, 1853, and A.trux Walker, 1853 under A.longus Walker, 1853.

Cryptochrysa Freitas Penny, a generic homonym, replaced by Titanochrysa Sosa Freitas (Neuroptera: Chrysopidae).

The green lacewing genus name Cryptochrysa Freitas Penny 2001 is identified as a junior homonym of Cryptochrysa Hampson 1926 (Lepidoptera: Noctuidae). Moreover, Titanochrysa Sosa Freitas 2012 is determined to be an available junior synonym of Cryptochrysa Freitas Penny. Thus, Titanochrysa becomes the substitute name for the preoccupied generic name. Here, we also provide (i) new information and images for Titanochrysa chloros (Freitas Penny) comb. nov., the only species ever included in the chrysopid genus Cryptochrysa, (ii) a clarified set of diagnostic features for the reconstituted genus Titanochrysa, and (iii) a key and images for identifying the six described Titanochrysa species.

The genus Vieira Navás (Neuroptera: Chrysopidae): a new species, a key to the species, and new geographic records.

Vieira Navás is a small genus of Neotropical green lacewings in the tribe Belonopterygini. Here, a fifth species, Vieira flinti sp. nov., is described from Venezuela and Brazil. This species shares significant features with disparate members of the genus; thus the genus as a whole is now known to express a more continuous range of morphological variation than previously recognized. We present a key for identifying Vieira species and new geographic records for previously described species in the genus.

Corrigenda: Revision of the Neotropical green lacewing genus Ungla (Neuroptera, Chrysopidae). ZooKeys 674: 1-188. https://doi.org/10.3897/zookeys.674.11435.

[This corrects the article DOI: 10.3897/zookeys.674.11435.].

Revision of the Neotropical green lacewing genus Ungla (Neuroptera, Chrysopidae).

Here, Ungla Navás, 1914, a poorly known Neotropical genus is reviewed. Twenty-five valid species are recognized; seven of them are new to science: Ungla adamsisp. n., U. elbergisp. n., U. grandispiraculasp. n., U. mexicanasp. n., U. pennyisp. n., U. quchapampasp. n., U. stangeisp. n.; and five are transferred to Ungla from other genera: U. bolivari (Banks), U. chacranella (Banks), U. siderocephala (Navás), U. steinbachi (Navás), and U. banksi Tauber, new replacement name. In addition, ten new synonymies are identified. For each species, a full nomenclatural history, diagnosis, description or redescription with images, literature citations, and available information on the distribution and biology are provided. Name-bearing types were examined for each species, and images of most are included. Keys based on external features are provided for species identifications. As a result of this study, three generalizations appear: (1) The genital morphology of both males and females of Ungla species is very conserved. All species express a common structural pattern, the components of which vary only slightly among species. (2) Ungla species appear to fall into two geographically distinct groups: about one third (n=7) of the species are recorded from southern South America (specifically Argentina and Brazil) and the other approximately two thirds of the species (n=18) from more northern regions of Neotropical America [Andean and Caribbean regions, Central America, and southern Mexico (Chiapas)]. None of the species from either of the regions is known to overlap into the other region. (3) Available information on the immature stages and natural history of species in Ungla is meagre.

Rediscovery of Nuvol umbrosus Navás (Neuroptera, Chrysopidae, Leucochrysini): a redescription and discussion.

The monotypic leucochrysine genus Nuvol was previously known from three specimens of Nuvol umbrosus Navás, collected in the Atlantic Forest region of Brazil. For many years these specimens have been missing, and the genus has remained without a modern description. Here, the species is redescribed based on two newly discovered specimens (females) from the Amazonian region. The female terminalia are relatively simple, except for the subgenitale, which is enlarged, folded into two sections, and heavily sclerotized. Unique aspects of the wing venation and the unusual pattern of banding on the wings support the retention of Nuvol as a valid genus within the Leucochrysini. There are differences between the Amazonian specimens studied here and the earlier descriptions based on specimens from the Atlantic Forest. These differences may indicate the presence of two distinct, geographically separated species within the genus. However, largely because we do not know the sexes of the earlier specimens, we are treating the differences discovered in the two female specimens as expressions of intraspecific variation.

Apochrysinae (Neuroptera: Chrysopidae): new larval description and subfamilial comparisons.

Although Apochrysinae is considered an important, ancient clade of Chrysopidae, its immature stages are very poorly known. Larvae are rarely seen, and descriptions are limited to the third instar of only one species in the genus Apochrysa. Herein, the larvae (all instars) of a second species, Apochrysa voeltzkowi (Weele), are described, and the features of the two Apochrysa species are compared with those of other chrysopids. The results provide support (one potential synapomorphy) for a phylogenetic relationship between Apochrysinae and Nothochrysinae and additional support for two synapomorphies previously proposed for Nothochrysinae. Diagnostic larval characteristics are proposed for the genus Apochrysa, and species-specific features are identified for the two known Apochrysa larvae.

Kymachrysa, a new genus of Nearctic Green Lacewings (Neuroptera, Chrysopidae, Chrysopini).

Two North American species of green lacewings have undergone a number of changes in their generic assignments and are currently classified as incertae sedis. Here we demonstrate that adults (both sexes) and larvae of these species share a set of features that distinguishes them from currently described genera. Thus, to promote nomenclatural stability in Chrysopidae, we describe Kymachrysa, a gen. n. that contains the two species - Kymachrysa intacta (Navás), comb. n. and Kymachrysa placita (Banks), comb. n.. Also, we present modifications for the current generic-level key, illustrations, as well as biological information for identifying the genus and its known species.

Two common and problematic leucochrysine species - Leucochrysa (Leucochrysa) varia (Schneider) and L. (L.) pretiosa (Banks) (Neuroptera, Chrysopidae): redescriptions and synonymies.

We dedicate this article to the memory of Sergio de Freitas, FCAV-UNESP, Jaboticabal, São Paulo, Brazil (deceased, 2012). He was an active and enthusiastic Neuropterist and the cherished mentor and friend of Francisco Sosa. Leucochrysa McLachlan is the largest genus in the Chrysopidae, yet it has received relatively little taxonomic attention. We treat two problematic and common Leucochrysa species - Leucochrysa (Leucochrysa) varia (Schneider, 1851) and Leucochrysa (Leucochrysa) pretiosa (Banks, 1910). Both are highly variable in coloration and were described before the systematic importance of chrysopid genitalia was recognized. Recent studies show that these species occur within a large complex of cryptic species and that they have accumulated a number of taxonomic problems. We identify new synonymies for each of the species-for Leucochrysa (Leucochrysa) varia: Leucochrysa (Leucochrysa) ampla (Walker, 1853), Leucochrysa internata (Walker, 1853), and Leucochrysa (Leucochrysa) walkerina Navás, 1913; for Leucochrysa (Leucochrysa) pretiosa: Leucochrysa (Leucochrysa) erminea Banks, 1946. The synonymy of Leucochrysa delicata Navás, 1925 with Leucochrysa (Leucochrysa) pretiosa is stabilized by the designation of a neotype. The following species, which were previously synonymized with Leucochrysa (Leucochrysa) varia or Leucochrysa (Leucochrysa) pretiosa, are reinstated as valid: Leucochrysa (Leucochrysa) phaeocephala Navás, 1929, Leucochrysa (Leucochrysa) angrandi (Navás, 1911), and Leucochrysa (Leucochrysa) variata (Navás, 1913). To help stabilize Leucochrysa taxonomy, lectotypes are designated for Allochrysa pretiosa and Allochrysa variata. Finally, Leucochrysa vegana Navás, 1917 is considered a nomen dubium.

ResumenLeucochrysa McLachlan es el género más grande de Chrysopidae, sin embargo, ha recibido poca atención taxonómica. Dos de las especies más comunes y problemáticas de Leucochrysa son estudiadas ­ Leucochrysa (Leucochrysa) varia (Schneider, 1851) y Leucochrysa (Leucochrysa) pretiosa (Banks, 1910). Ambas son altamente variables en coloración y fueron descritas antes de que se reconociera la importancia de la genitalia en la sistemática de Chrysopidae. Estudios recientes demuestran que estas especies pertenecen a un complejo de especies crípticas, razón por la cual han acumulado numerosos problemas taxonómicos. En este estudio se identificaron las siguientes sinonimias; para Leucochrysa (Leucochrysa) varia: Leucochrysa (Leucochrysa) ampla (Walker, 1853), Leucochrysa internata (Walker, 1853), y Leucochrysa (Leucochrysa) walkerina Navás, 1913; para Leucochrysa (Leucochrysa) pretiosa: Leucochrysa (Leucochrysa) erminea Banks, 1946. La sinonimia de Leucochrysa delicata Navás, 1925 con Leucochrysa (Leucochrysa) pretiosa es estabilizada por la designación de un neotipo. Las especies Leucochrysa (Leucochrysa) phaeocephala Navás, 1929, Leucochrysa (Leucochrysa) angrandi (Navás, 1911) y Leucochrysa (Leucochrysa) variata (Navás, 1913) las cuales fueron previamente sinonimizadas con Leucochrysa (Leucochrysa) varia o Leucochrysa (Leucochrysa) pretiosa, son reestablecidas como válidas. Para ayudar a estabilizar la taxonomía de Leucochrysa, se designaron lectotipos para Allochrysa pretiosa y Allochrysa variata. Finalmente, Leucochrysa vegana Navás, 1917 es considerada como un nomen dubium.

ResumoLeucochrysa McLachlan é o maior gênero de Chrysopidae; no entanto, ele tem recebido relativamente pouca atenção taxonômica. Nesse trabalho, duas espécies de Leucochrysa problemáticas e comuns são analisadas - Leucochrysa (Leucochrysa) varia (Schneider, 1851) e Leucochrysa (Leucochrysa) pretiosa (Banks, 1910). Ambas foram descritas antes da importância sistemática da genitália de crisopídeos ter sido reconhecida e são altamente variáveis na coloração. Estudos recentes têm demonstrado que essas espécies ocorrem dentro de um grande complexo de espécies crípticas e que elas têm acumulado diversos problemas taxonômicos. Foram identificadas novas sinonímias para cada uma das espécies - para Leucochrysa (Leucochrysa) varia: Leucochrysa (Leucochrysa) ampla (Walker, 1853), Leucochrysa internata (Walker, 1853) e Leucochrysa (Leucochrysa) walkerina Navás, 1913; para Leucochrysa (Leucochrysa) pretiosa: Leucochrysa (Leucochrysa) erminea Banks, 1946. A sinonímia de Leucochrysa delicata Navás, 1925 com Leucochrysa (Leucochrysa) pretiosa é estabilizada pela designação de um neótipo. As seguintes espécies, que eram previamente sinonimizadas com Leucochrysa (Leucochrysa) varia ou Leucochrysa (Leucochrysa) pretiosa, são reintegradas como válidas: Leucochrysa (Leucochrysa) phaeocephala Navás, 1929, Leucochrysa (Leucochrysa) angrandi (Navás, 1911) e Leucochrysa (Leucochrysa) variata (Navás, 1913). Para auxiliar na estabilização da taxonomia de Leucochrysa, lectótipos são designados para Allochrysa pretiosa e Allochrysa variata. Finalmente, Leucochrysa vegana Navás, 1917 é considerada nomen dubium.

Larvae of five horticulturally important species of Chrysopodes (Neuroptera, Chrysopidae): shared generic features, descriptions and keys.

An expanded list of generic level larval characteristics is presented for Chrysopodes; it includes a reinterpretation of the mesothoracic and metathoracic structure and setation. Keys, descriptions and images of Semaphoront A (first instar) and Semaphoront B (second and third instars) are offered for identifying five species of Chrysopodes (Chrysopodes) that are commonly reported from horticultural habitats in the Neotropical region.

ResumoUma lista expandida das características larvais em nível de gênero é apresentada para Chrysopodes, incluindo a reinterpretação da estrutura e das cerdas do mesotórax e metatórax. Chaves, descrições e imagens do semaforonte A (primeiro instar) e semaforonte B (segundo e terceiro instares) são fornecidas para a identificação de cinco espécies de Chrysopodes (Chrysopodes) comumente encontradas em habitats hortícolas na região Neotropical.

Santocellus (Neuroptera, Chrysopidae, Leucochrysini): taxonomic changes, new description, and a key to the species.

Santocellus is a small Neotropical genus of leucochrysine lacewings that only recently was separated from Leucochrysa. Here, the features of the Leucochrysa risi Esben-Petersen holotype (a female) are described and shown to support the species' transfer to Santocellus and the continued retention of the genus Santocellus as separate from Leucochrysa. The valid name for the species becomes Santocellus risi (Esben-Petersen, 1933), comb. n., and Santocellus bullata (Tauber, 2007) is recognized as a syn. n. of Santocellus risi. Currently, this species is reported only from Peru. An illustrated key is provided for distinguishing the known species in the genus Santocellus.