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).
The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.
Myrmeleontiformia are an ancient group of lacewing insects characterized by predatory larvae with unusual morphologies and behaviours. Mostly soil dwellers with a soft cuticle, their larvae fossilize only as amber inclusions, and thus their fossil record is remarkably sparse. Here, we document a disparate assemblage of myrmeleontiform larvae from the mid-Cretaceous amber (99 Ma) of Myanmar, evidence of a considerable diversification. Our cladistic analysis integrating extant and extinct taxa resolves the fossils as both stem- and crown-groups. Similarities between extinct and extant species permit inferences of larval ethology of the fossil species through statistical correlation analyses with high support, implying that morphological disparity matched behavioural diversity. An improved understanding of the evolutionary history of antlions and relatives supports the conclusion that hunting strategies, such as camouflage and fossoriality, were acquired early within the lineage.
The larvae of the European Myrmeleontidae are reviewed with the aim to ease their identification, covering 15 genera and 28 species. Diagnostic characters and illustrations are given for each taxon. Larvae of the genera Nemoleon and Macronemurus are described for the first time while Megistopus, Neuroleon and Myrmeleon are revised. The larvae of Dendroleon pantherinus (Fabricius), Macronemurus appendiculatus (Latreille), Megistopus lucasi (Navás), Nemoleon notatus (Rambur), Neuroleon arenarius (Navás), Neuroleon assimilis (Navás), Neuroleon nemausiensis (Borkhausen), Cueta lineosa (Rambur) and Myrmeleon gerlindae (Hölzel) are described or accurately depicted for the first time.
A phylogenetic analysis of selected oestroid taxa based on 66 morphological traits and sequences from three nuclear protein‐coding genes (CAD, MAC, MCS) resolved the composition and phylogenetic position of the former subfamily Polleniinae of the Calliphoridae – here resurrected at family rank as Polleniidae Brauer & Bergenstamm, 1889 stat. rev. Six species are transferred from the family Rhinophoridae to the Polleniidae: the Palaearctic genus Alvamaja Rognes, along with its single species Alvamaja chlorometallica Rognes, and five Afrotropical species comprising the carinata‐group formerly in the genus Phyto Robineau‐Desvoidy but here assigned to genus Morinia Robineau‐Desvoidy, i.e. M. carinata (Pape, 1987) comb.n., M. lactineala (Pape, 1997) comb.n., M. longirostris (Crosskey, 1977) comb.n., M. royi (Pape, 1997) comb.n. and M. stuckenbergi (Crosskey, 1977) comb.n. The Polleniidae are monophyletic and, in agreement with most recent phylogenetic reconstructions, sister to the Tachinidae. The female of A. chlorometallica and a new species of Morinia of the carinata‐group (Morinia tsitsikamma sp.n. from South Africa) are described. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:25B0C220-DEE4-4B0C-88EA-35FDE298EBC5.
Fossils sometimes show unusual morphological features absent in living organisms, making it difficult to reconstruct both their affinity and their function. We describe here a new lacewing larva, Ankyloleon caudatus gen. et sp.n. (Neuroptera) from the Cretaceous amber of Myanmar, characterized by an abdomen unique among insects, with ‘tail‐like’ terminal segments bearing a ventral pair of vesicles. Phase‐contrast X‐ray microtomography reveals that these structures were dense and equipped with a median duct, suggesting that they were likely pygopods used for locomotion, holding the position through adhesive secretions. Our phylogenetic analyses, combining genomic and morphological data from both living and fossil lacewings, proved critical to placing Ankyloleon gen.n. on the lacewing tree of life as an early representative of the antlion clade, Myrmeleontiformia. These results corroborate the view that derived myrmeleontiform lacewings ‘experimented’ with unusual combinations of features and specializations during their evolutionary history, some of which are now lost. Zoobank registration: urn:lsid:zoobank.org:pub:0C0AC565‐1AC9‐42CC‐831D‐EDA38BA36F64
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
The world Rhinophoridae are catalogued, recognising 33 genera and 177 species. Nomenclatural information is provided for all genus-group and species-group names, including lists of synonyms and name-bearing type data. Species distributions are recorded by country. A key to the world genera is presented. Four new genera are erected to accommodate five new species, which do not fit within any of the current generic concepts in Rhinophoridae, according to the results of a morphology-based phylogenetic analysis: Marshallicona Cerretti & Pape with type species Marshallicona quitu Cerretti & Pape, gen. et sp. nov. (Ecuador); Maurhinophora Cerretti & Pape with type species Maurhinophora indoceanica Cerretti & Pape, gen. et sp. nov. (Mauritius); Neotarsina Cerretti & Pape with type species Neotarsina caraibica Cerretti & Pape, gen. et sp. nov. (Trinidad and Tobago) and Neotarsina andina Cerretti & Pape, sp. nov. (Peru); Kinabalumyia Cerretti & Pape with type species Kinabalumyia pinax Cerretti & Pape, gen. et sp. nov. (Malaysia, Sabah). The genus Aporeomyia Pape & Shima (type species Aporeomyia antennalis Pape & Shima), originally assigned to Tachinidae, is here reassigned to Rhinophoridae based on a reassessment of the homologies of the male terminalia. The following five species-group names, which were previously treated as junior synonyms or nomina dubia, are recognised as valid species names: Acompomintho caucasica (Villeneuve, 1908), stat. rev. [from nomen dubium to valid species]; Acompomintho sinensis (Villeneuve, 1936), stat. rev. [from nomen dubium to valid species]; Stevenia bertei (Rondani, 1865), stat. rev. [from nomen dubium to valid species]; Stevenia sardoa Villeneuve, 1920, stat. rev. [from junior synonym of Rhinophora deceptoria Loew, 1847 to valid species]; Stevenia subalbida (Villeneuve, 1911), stat. rev. [from junior synonym of Rhinophora deceptoria Loew, 1847 to valid species]. Reversal of precedence is invoked for the following case of subjective synonymy to promote stability in nomenclature: Rhinophora lepida (Meigen, 1824), nomen protectum, and Musca parcus Harris, 1780: 144, nomen oblitum. New generic and specific synonymies are proposed for the following two names: Mimodexia Rohdendorf, 1935, junior synonym of Tromodesia Rondani, 1856, syn. nov. and Ptilocheta tacchetti Rondani, 1865, junior synonym of Stevenia obscuripennis (Loew, 1847), syn. nov. The following new combinations are proposed: Acompomintho sinensis (Villeneuve, 1936), comb. nov. [transferred from Tricogena Robineau-Desvoidy, 1830]; Tromodesia guzari (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia intermedia (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia lindneriana (Rohdendorf, 1961), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia magnifica (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia obscurior (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia pallidissima (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia setiventris (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935] and Tromodesia shachrudi (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935].
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