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.

Attraction of the Green Lacewing Chrysoperla comanche (Neuroptera: Chrysopidae) to Yeast.

Many adult Chrysoperla comanche (Stephens) green lacewings were caught in traps baited with live yeast cultures during tests designed to catch olive fruit flies. All 13 yeast species tested were more attractive than the industry-standard dried torula yeast (Cyberlindnera jadinii; syn. Candida utilis). Live C. jadinii culture attracted significantly more lacewings than the inactive dried-pellet form of the same yeast species, demonstrating that volatiles from live yeast cultures attract adults of this lacewing. Odor profiles for two of the highly active yeasts tested herein (Lachancea thermotolerans and Solicoccozyma terrea) were similar to that for Metschnikowia pulcherrima, a yeast species isolated earlier from the gut diverticulum of Chrysoperla rufilabris. A new Metschnikowia species (M. chrysoperlae), along with two new Candida spp. that were recently realigned to one of the Metschnikowia clades (M. picachoensis and M. pimensis), were also identified from the diverticulum of C. comanche. Thus, one clade of Metschnikowia yeasts that commonly occur in floral nectar appears to exhibit mutualistic symbioses with Chrysoperla green lacewings. Both male and female C. comanche adults were attracted in the present study, and we speculate that males have exploited this symbiosis by offering Metschnikowia-laden regurgitant, including attractive volatiles, to females ('mating trophallaxis') as a nuptial gift.

Phylogeny and Evolution of Neuropterida: Where Have Wings of Lace Taken Us?

The last 25 years of phylogenetic investigation into the three orders constituting the superorder Neuropterida-Raphidioptera, Megaloptera, and Neuroptera-have brought about a dramatic revision in our understanding of the evolution of lacewings, snakeflies, dobsonflies, and their diverse relatives. Phylogenetic estimations based on combined analyses of diverse data sources, ranging from adult and larval morphology to full mitochondrial genomic DNA, have begun to converge on similar patterns, many times in accordance with hypotheses put forth by Cyril Withycombe nearly a century ago. These data, in combination with information from the fossil record, have given a revised perspective on the historical evolution and classification of Neuropterida, necessitating an overhaul of their organization and providing focus and insight on fruitful future efforts for neuropterology.

Anchored phylogenomics unravels the evolution of spider flies (Diptera, Acroceridae) and reveals discordance between nucleotides and amino acids.

The onset of phylogenomics has contributed to the resolution of numerous challenging evolutionary questions while offering new perspectives regarding biodiversity. However, in some instances, analyses of large genomic datasets can also result in conflicting estimates of phylogeny. Here, we present the first phylogenomic scale study of a dipteran parasitoid family, built upon anchored hybrid enrichment and transcriptomic data of 240 loci of 43 ingroup acrocerid taxa. A new hypothesis for the timing of spider fly evolution is proposed, wielding recent advances in divergence time dating, including the fossilized birth-death process to show that the origin of Acroceridae is younger than previously proposed. To test the robustness of our phylogenetic inferences, we analyzed our datasets using different phylogenetic estimation criteria, including supermatrix and coalescent-based approaches, maximum-likelihood and Bayesian methods, combined with other approaches such as permutations of the data, homogeneous versus heterogeneous models, and alternative data and taxon sets. Resulting topologies based on amino acids and nucleotides are both strongly supported but critically discordant, primarily in terms of the monophyly of Panopinae. Conflict was not resolved by controlling for compositional heterogeneity and saturation in third codon positions, which highlights the need for a better understanding of how different biases affect different data sources. In our study, results based on nucleotides were both more robust to alterations of the data and different analytical methods and more compatible with our current understanding of acrocerid morphology and patterns of host usage.

Mitochondrial phylogenomics illuminates the evolutionary history of Neuropterida.

Neuroptera (lacewings) and allied orders Megaloptera (dobsonflies, alderflies) and Raphidioptera (snakeflies) are predatory insects and together make up the clade Neuropterida. The higher-level relationships within Neuropterida have historically been widely disputed with multiple competing hypotheses. Moreover, the evolution of important biological innovations among various Neuropterida families, such as the origin, timing and direction of transitions between aquatic and terrestrial habitats of larvae, remains poorly understood. To investigate the origin and diversification of lacewings and their allies, we undertook phylogenetic analyses of mitochondrial genomes of all families of Neuropterida using Bayesian inference, maximum likelihood and maximum parsimony methods. We present a robust, fully resolved phylogeny and divergence time estimation for Neuropterida with strong statistical support for almost all nodes. Mitochondrial sequence data are typified by significant compositional heterogeneity across lineages, and parsimony and models assuming homogeneous rates did not recover Neuroptera as monophyletic. Only a model accounting for compositional heterogeneity (i.e. CAT-GTR) recovered all orders of Neuropterida as monophyletic. Significant findings of the mitogenomic phylogeny include recovering Raphidioptera as sister to Megaloptera plus Neuroptera. The sister family of all other lacewings are the dusty-wings (Coniopterygidae), rather than Nevrorthidae. Nevrorthidae are instead returned to their traditional position as the sister group of the spongilla-flies (Sisyridae) and closely related to Osmylidae. Our divergence time analysis indicates that the Mesozoic was indeed a 'golden age' for lacewings, with most families of Neuropterida diverging during the Triassic and Jurassic and all extant families present by the Early Cretaceous. Based on ancestral character state reconstructions of larval habitat we evaluate competing hypotheses regarding the life style of early neuropteridan larvae as either aquatic or terrestrial.

The phylogeny of brown lacewings (Neuroptera: Hemerobiidae) reveals multiple reductions in wing venation.

BACKGROUND: The last time the phylogenetic relationships among members of the family Hemerobiidae were studied quantitatively was over 12 years ago and based exclusively on morphology. Our study builds upon this morphological evidence by adding sequence data from three gene loci to provide a total evidence phylogeny of brown lacewings (Neuroptera: Hemerobiidae). Thirty-seven species representing nineteen Hemerobiidae genera were compared with outgroups from the families Ithonidae, Psychopsidae and Chrysopidae in Bayesian and parsimony analyses using a single nuclear gene (CAD) and two mitochondrial (16S rDNA and Cytochrome Oxidase I) genes. We compare divergence time estimates of Hemerobiidae cladogenesis under the two most commonly used relaxed clock models and discuss the evolution of wing venation in the family. RESULTS: We recovered a phylogeny largely incongruent with previously published morphological studies, although all but two subfamilies (i.e., Notiobiellinae and Drepanacrinae) were recovered as monophyletic. We found the subfamily Drepanacrinae paraphyletic with respect to Psychobiellinae, and Notiobiellinae to be polyphyletic. We thus offer a revised concept of Notiobiellinae, comprising only Notiobiella Banks, and erect a new subfamily Zachobiellinae including the remaining genera previously placed in Notiobiellinae. Psychobiellinae is synonymized with Drepanacrinae. Unlike the previous hypothesis that proposed a remarkably laddered topology, our tree suggests that hemerobiids diverged as three main clades. Moreover, in contrast to the vein proliferation hypothesis, we found that hemerobiids have instead undergone multiple reductions in the number of radial veins, this scenario questions the relevance of this character as diagnostic of various subfamilies CONCLUSIONS: Our phylogenetic hypothesis and divergence times analysis suggest that extant hemerobiids originated around the end of the Triassic and evolved as three distinct clades that diverged from one another during the Late Jurassic to Early Cretaceous. Contrary to earlier phylogenetic hypotheses, Carobius Banks (Carobiinae) is sister to the previously unplaced genus Notherobius New in a clade more closely related to Sympherobiinae, Megalominae and Zachobiellinae subfam. nov. The addition of taxa which are not available for DNA sequencing should be the focus of future studies, especially Adelphohemerobius Oswald, which is particularly important to test our inferences regarding the evolution of wing venation in Hemerobiidae.

Phylogeny of split-footed lacewings (Neuroptera, Nymphidae), with descriptions of new Cretaceous fossil species from China.

A phylogeny of the lacewing family Nymphidae based on morphology and DNA sequences is presented including representatives of all living genera and selected fossil genera. Widely distributed Jurassic and Cretaceous genera gave rise to recent taxa now restricted to Australasia. Two previously defined clades (i.e. Nymphinae and Myiodactylinae) were recovered and reflect the diverging adult and larval morphology of members of these two subfamilies. From Chinese Cretaceous deposits, a new genus (Spilonymphes gen. nov.) is described with one new species, as well as new species described in the genera Baissoleon Makarkin and Sialium Westwood.

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.

Revision of the South American stiletto fly genera Entesia Oldroyd and Melanothereva Malloch, with the description of a new genus (Therevidae: Agapophytinae).

The South American members of the stiletto fly subfamily Agapophytinae (Diptera: Therevidae) occur in the South American Transition Zone, predominantly in Chile and southwestern Argentina. The four agapophytine genera described from this region are Entesia Oldroyd, 1968; Melanothereva Malloch 1932; Pachyrrhiza Philippi, 1865; and more recently, Sigalopella Irwin & Winterton, 2020. Here we revise Melanothereva and Entesia and describe the new monotypic genus Pachylopella. Keys to South American Agapophytinae genera and species of Melanothereva and Entesia are provided.

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.

A new species of Amplisegmentum Webb (Diptera: Therevidae) from Venezuela.

A new species of Amplisegmentum Webb (Therevidae: Therevinae) is described from Venezuela, a previously monotypic genus otherwise known only from Ecuador. A revised diagnosis of the genus and a key to species is presented.

Revision of South American stiletto fly genus Argolepida Metz amp; Irwin (Diptera: Therevidae: Therevinae).

The stiletto fly genus Argolepida Metz Irwin, 2003 is revised. Members of this genus are diminutive, yet distinctive, flies restricted to southern South America. Argolepida rivulosa (Krber, 1928) is redescribed and seven new species are described and figured. A key to species is presented.

Revision of the Patagonian stiletto fly genus Pachyrrhiza Philippi (Therevidae: Agapophytinae).

In South America, the stiletto fly subfamily Agapophytinae (Diptera: Therevidae) occurs predominantly in the Patagonian region where it is represented by four genera: Pachyrrhiza Philippi, Melanothereva Malloch, Entesia Oldroyd and Sigalopella Irwin Winterton. Here we revise the genus Pachyrrhiza with an additional four new species described and figured.

A new species of Joguina Navás, 1912 from India (Neuroptera: Chrysopidae).

A new species of the delicate lacewing, Joguina Navás, 1912 (Chrysopidae: Apochrysinae) is described and figured from south-western India. A key to species of Joguina is presented as well as revised diagnoses of both Joguina and its sister genus, Lainius Navás, 1913 stat. rev.

Single-copy nuclear genes resolve the phylogeny of the holometabolous insects.

BACKGROUND: Evolutionary relationships among the 11 extant orders of insects that undergo complete metamorphosis, called Holometabola, remain either unresolved or contentious, but are extremely important as a context for accurate comparative biology of insect model organisms. The most phylogenetically enigmatic holometabolan insects are Strepsiptera or twisted wing parasites, whose evolutionary relationship to any other insect order is unconfirmed. They have been controversially proposed as the closest relatives of the flies, based on rDNA, and a possible homeotic transformation in the common ancestor of both groups that would make the reduced forewings of Strepsiptera homologous to the reduced hindwings of Diptera. Here we present evidence from nucleotide sequences of six single-copy nuclear protein coding genes used to reconstruct phylogenetic relationships and estimate evolutionary divergence times for all holometabolan orders. RESULTS: Our results strongly support Hymenoptera as the earliest branching holometabolan lineage, the monophyly of the extant orders, including the fleas, and traditionally recognized groupings of Neuropteroidea and Mecopterida. Most significantly, we find strong support for a close relationship between Coleoptera (beetles) and Strepsiptera, a previously proposed, but analytically controversial relationship. Exploratory analyses reveal that this relationship cannot be explained by long-branch attraction or other systematic biases. Bayesian divergence times analysis, with reference to specific fossil constraints, places the origin of Holometabola in the Carboniferous (355 Ma), a date significantly older than previous paleontological and morphological phylogenetic reconstructions. The origin and diversification of most extant insect orders began in the Triassic, but flourished in the Jurassic, with multiple adaptive radiations producing the astounding diversity of insect species for which these groups are so well known. CONCLUSION: These findings provide the most complete evolutionary framework for future comparative studies on holometabolous model organisms and contribute strong evidence for the resolution of the 'Strepsiptera problem', a long-standing and hotly debated issue in insect phylogenetics.

A new bee-mimicking stiletto fly (Therevidae) from China discovered on iNaturalist.

The Chinese stiletto fly fauna is poorly known, with few species previously described and no endemic genera. A new genus and species of charismatic stiletto fly is described from northern China. Although the species was previously known from a poorly preserved specimen, the true form of this apparent bumble bee-mimic was not known until images of a live individual were discovered amongst photographs posted on iNaturalist, an online citizen scientist and biodiversity social network.

Review of the green lacewing genus Apochrysa Schneider (Neuroptera: Chrysopidae).

Delicate green lacewings in the genus Apochrysa Schneider (Neuroptera: Chrysopidae: Apochrysinae) are reviewed with each species diagnosed and figured; a key to species is included.

A new genus of therevine stiletto flies from South America (Diptera: Therevidae).

A new genus of Therevidae (Dasythereva gen. n.) is described from Argentina and Chile and comprises two distinctive new species (Dasythereva penai sp. n., D. patagonia sp. n.) from sandy habitats. A key to genera of Neotropical Therevinae is included to accommodate the inclusion of this new genus.

A new stiletto fly genus from South America (Diptera: Therevidae: Agapophytinae).

The South American members of the stiletto fly subfamily Agapophytinae (Diptera: Therevidae) comprise three genera that occur predominantly in the western region. Here we describe a newly discovered genus, Sigalopella gen. n., from Chile containing four new species.

New stiletto flies from New Caledonia (Therevidae, Agapophytinae).

Stiletto-flies (Diptera: Therevidae) are highly diverse and species-rich in Australia and New Zealand, yet relatively few species have been recorded from neighbouring Papua New Guinea, Indonesia and throughout the remainder of Oceania. Indeed, in New Caledonia only a single species of the widely distributed Australasian genus Anabarhynchus Macquart (Therevinae) is previously known. Herein we describe two new agapophytine genera (i.e., Jeanchazeauia gen. nov., Calophytus gen. nov.), together comprising nine charismatic new species; this represents a first record of the subfamily from New Caledonia. The new genera and species are described and figured.