Analysis of anchored hybrid enrichment (AHE) data under a variety of analytical parameters for a broadly representative sample of taxa (136 species representing all extant families) recovered a well-resolved and strongly supported tree for the higher phylogeny of Neuropterida that is highly concordant with previous estimates based on DNA sequence data. Important conclusions include: Megaloptera is sister to Neuroptera; Coniopterygidae is sister to all other lacewings; Osmylidae, Nevrorthidae and Sisyridae are recovered as a monophyletic Osmyloidea, and Rhachiberothidae and Berothidae were recovered within a paraphyletic Mantispidae. Contrary to previous studies, Chrysopidae and Hemerobiidae were not recovered as sister families and morphological similarities between larvae of both families supporting this assumption are reinterpreted as symplesiomorphies. Relationships among myrmeleontoid families are similar to recent studies except Ithonidae are placed as sister to Nymphidae. Notably, Ascalaphidae render Myrmeleontidae paraphyletic, again calling into question the status of Ascalaphidae as a separate family. Using statistical binning of partitioned loci based on a branch-length proxy, we found that the diversity of phylogenetic signal across partitions was minimal from the slowest to the fastest evolving loci and varied little over time. Ancestral character-state reconstruction of the sclerotization of the gular region in the larval head found that although it is present in Coleoptera, Raphidioptera and Megaloptera, it is lost early in lacewing evolution and then regained twice as a nonhomologous gula-like sclerite in distantly related clades. Reconstruction of the ancestral larval habitat also indicates that the ancestral neuropteridan larva was aquatic, regardless of the assumed condition (i.e., aquatic or terrestrial) of the outgroup (Coleopterida).

No abstract

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.

The wings of insects are one of their most prominent features and embody numerous characters and modifications congruent with the variety of their lifestyles. However, despite their evolutionary relevance, homology statements and nomenclature of wing structures remain understudied and sometimes confusing. Early studies on wing venation homologies often assumed Neuropterida (the superorder comprising the orders Raphidioptera, Megaloptera, and Neuroptera: snakeflies, alderflies and dobsonflies, and lacewings) to be ancient among Pterygota, and therefore relied on their pattern of venation for determining groundplans for insect wing venation schemata and those assumptions reciprocally influenced the interpretation of lacewing wings. However, Neuropterida are in fact derived among flying insects and thus a reconsideration of their wings is crucial. The identification of the actual wing venation of Neuropterida is rendered difficult by fusions and losses, but these features provide systematic and taxonomically informative characters for the classification of the different clades within the group. In the present study, we review the homology statements of wing venation among Neuropterida, with an emphasis on Chrysopidae (green lacewings), the family in which the highest degree of vein fusion is manifest. The wing venation of each order is reviewed according to tracheation, and colored schemata of the actual wing venation are provided as well as detailed illustrations of the tracheation in select families. According to the results of our study of vein tracheation, new homology statements and a revised nomenclature for veins and cells are proposed.

We present a time-calibrated phylogeny of the charismatic green lacewings (Neuroptera: Chrysopidae). Previous phylogenetic studies on the family using DNA sequences have suffered from sparse taxon sampling and/or limited amounts of data. Here we combine all available previously published DNA sequence data and add to it new DNA sequences generated for this study. We analysed these data in a supermatrix using Bayesian and maximum likelihood methods and provide a phylogenetic hypothesis for the family that recovers strong support for the monophyly of all subfamilies and resolves relationships among a large proportion of chrysopine genera. Chrysopinae tribes Leucochrysini and Belonopterygini were recovered as monophyletic sister clades, while the species-rich tribe Chrysopini was rendered paraphyletic by Ankylopterygini. Relationships among the subfamilies were resolved, although with relatively low statistical support, and the topology varied based on the method of analysis. Greatest support was found for Apochrysinae as sister to Nothochrysinae and Chrysopinae, which is in contrast to traditional concepts that place Nothochrysinae as sister to the rest of the family. Divergence estimates suggest that the stem groups to the various subfamilies diverged during the Triassic-Jurassic, and that stem groups of the chrysopine tribes diverged during the Cretaceous.

The Red Queen hypothesis (RQH) is both familiar and murky, with a scope and range that has broadened beyond its original focus. Although originally developed in the palaeontological arena, it now encompasses many evolutionary theories that champion biotic interactions as significant mechanisms for evolutionary change. As such it de-emphasizes the important role of abiotic drivers in evolution, even though such a role is frequently posited to be pivotal. Concomitant with this shift in focus, several studies challenged the validity of the RQH and downplayed its propriety. Herein, we examine in detail the assumptions that underpin the RQH in the hopes of furthering conceptual understanding and promoting appropriate application of the hypothesis. We identify issues and inconsistencies with the assumptions of the RQH, and propose a redefinition where the Red Queen's reign is restricted to certain types of biotic interactions and evolutionary patterns occurring at the population level.

Insects exhibit a wide diversity of anatomical specializations in their adult and immature stages associated with particular aspects of their biology. The order Neuroptera (lacewings, antlions, and their relatives) are a moderately diverse lineage of principally predatory animals, at least in their immature stages, as all have a modified piercing-sucking mandible-maxillary complex that allows them to drain fluids from their prey. As such, the larvae of various groups have evolved unique anatomical and behavioral specializations for approaching and subduing their prey, particularly the green lacewings (Chrysopidae), where immatures are also adept at camouflage [1-4]. Here we report the discovery of a unique mode of life among mid-Cretaceous mesochrysopids, an early stem group to modern green lacewings [5-7] exhibiting a combination of morphological modifications in both adults and larvae unknown among living and fossil Neuroptera, even across winged insects. The new mesochrysopids exhibit a uniquely prolonged thorax, elongate legs, and dramatically reduced hind wings in adults, and larvae have extremely elongate, slender legs with pectinate pretarsal claws and lacking trumpet-shaped empodia. The peculiarities of the larvae include features principally found in spider-associated insect groups, implying that these lacewings were early specialists on web-spinning spiders, either as active predators or kleptoparasites. This reveals a dramatic and ancient degree of ecological refinement in a major lineage of insect predators, for a food resource otherwise not utilized by most lacewings.

A phylogeny of green lacewings (Neuroptera: Chrysopidae) using anchored hybrid enrichment data is presented. Using this phylogenomic approach, we analysed 137 kb of sequence data (with < 10% missing) for 82 species in 50 genera of Chrysopidae under Bayesian and maximum likelihood criteria. We recovered a strongly supported tree topologically congruent with recently published phylogenies, especially relationships amongst higher-level groups. The subfamily Nothochrysinae was recovered as paraphyletic, with one clade sister to the rest of Chrysopidae, and the second clade containing the nominal genus (Nothochrysa Navás) as sister to the subfamily Apochrysinae. Chrysopinae was recovered as a monophyletic with the monobasic Nothancylini tribe n. sister to the rest of the subfamily. Leucochrysini was recovered sister to Belonopterygini, and Chrysopini was rendered paraphyletic with respect to Ankylopterygini. Divergence times and diversification estimates indicate a major shift in rate in ancestral Chrysopini at the end of the Cretaceous, and the extensive radiation of Chrysopinae, the numerically dominant clade of green lacewings, began in the Mid-Paleogene (c. 45 Ma). 514