New records and new synonyms of antlions (Neuroptera, Myrmeleontidae) from China

The antlions from the subfamilies Nemoleontinae and Myrmecaelurinae deposited in the Entomological Museum of the China Agricultural University (CAU) belong to 16 species, among which 11 are recorded for China for the first time. Four species with mostly Oriental distribution are recorded for the first time from the northern localities in Palaearctic China. The Dzungarian faunistic center of antlions with two species recently described from Turan is distinguished. Some species characteristic of Mongolia were found in adjacent China. Occurrence in China of three species, collected by Russian expeditions 100 years ago, is confirmed. Five species described by Yang from China have been synonymized with widely distributed species.     

Chromosome numbers in antlions (Myrmeleontidae) and owlflies (Ascalaphidae) (Insecta,Neuroptera)

A short review of main cytogenetic features of insects belonging to the sister neuropteran families Myrmeleontidae (antlions) and Ascalaphidae (owlflies) is presented, with a particular focus on their chromosome numbers and sex chromosome systems. Diploid male chromosome numbers are listed for 37 species, 21 genera from 9 subfamilies of the antlions as well as for seven species and five genera of the owlfly subfamily Ascalaphinae. The list includes data on five species whose karyotypes were studied in the present work. It is shown here that antlions and owlflies share a simple sex chromosome system XY/XX; a similar range of chromosome numbers, 2n = 14-26 and 2n = 18-22 respectively; and a peculiar distant pairing of sex chromosomes in male meiosis. Usually the karyotype is particularly stable within a genus but there are some exceptions in both families (in the genera Palpares and Libelloides respectively). The Myrmeleontidae and Ascalaphidae differ in their modal chromosome numbers. Most antlions exhibit 2n = 14 and 16, and Palparinae are the only subfamily characterized by higher numbers, 2n = 22, 24, and 26. The higher numbers, 2n = 20 and 22, are also found in owlflies. Since the Palparinae represent a basal phylogenetic lineage of the Myrmeleontidae, it is hypothesized that higher chromosome numbers are ancestral for antlions and were inherited from the common ancestor of Myrmeleontidae + Ascalaphidae. They were preserved in the Palparinae (Myrmeleontidae), but changed via chromosomal fusions toward lower numbers in other subfamilies.     

Total‐evidence phylogeny of the owlflies (Neuroptera,Ascalaphidae) supports a new higher‐level classification

The first large‐scale, total‐evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) is presented. A combined morphological and molecular dataset was analysed under several analytical regimes for 76 exemplars of Myrmeleontiformia (Psychopsidae, Nymphidae, Nemopteridae, Myrmeleontidae, Ascalaphidae), including 57 of Ascalaphidae. At the subordinal level, the families were recovered in all analyses in the form Psychopsidae + (Nymphidae + (Nemopteridae + (Myrmeleontidae + Ascalaphidae). In the DNA‐only maximum‐likelihood analysis, Ascalaphidae were recovered as paraphyletic with respect to the Myrmeleontidae and the tribe Ululodini. In both the parsimony and Bayesian total‐evidence analyses, however, the latter with strong support, traditional Ascalaphidae were recovered as monophyletic, and in the latter, Stilbopteryginae were placed as the immediate sister group. The long‐standing subfamilies Haplogleniinae and Ascalaphinae were not recovered as monophyletic in any analysis, nor were several of the included tribes of non‐ululodine Ascalaphinae. The Ululodini were monophyletic and well supported in all analyses, as were the New World Haplogleniinae and, separately, the African/Malagasy Haplogleniinae. The remaining Ascalaphidae, collectively, were also consistently cohesive, but included a genus that until now has been placed in the Haplogleniinae, Protidricerus. Protidricerus was discovered to express a well‐developed pleurostoma, a feature previously only encountered in divided‐eye owlflies. The feature traditionally used to differentiate the Haplogleniinae and Ascalaphinae, the entire or divided eye, can no longer be regarded as a spot‐diagnostic synapomorphy to separate these groups within the family. A new subfamilial classification based on these results is proposed and includes the following five subfamilies: Albardiinae, Ululodinae, Haplogleniinae, Melambrotinae and Ascalaphinae. In addition, the monophyletic containing group (Myrmeleontidae + (Palparidae + (Stilbopterygidae + Ascalaphidae))) is elevated to the rank of superfamily, as Myrmeleontoidea, in order to accommodate much‐needed taxonomic and nomenclatural restructuring anticipated to occur within the Ascalaphidae in the future. A list of genera included in each subfamily of Ascalaphidae is provided.     

Evolution of green lacewings (Neuroptera: Chrysopidae): an anchored phylogenomics approach

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).     

Specialized learning in antlions (Neuroptera: Myrmeleontidae), pit-digging predators, shortens vulnerable larval stage

Unique in the insect world for their extremely sedentary predatory behavior, pit-dwelling larval antlions dig pits, and then sit at the bottom and wait, sometimes for months, for prey to fall inside. This sedentary predation strategy, combined with their seemingly innate ability to detect approaching prey, make antlions unlikely candidates for learning. That is, although scientists have demonstrated that many species of insects possess the capacity to learn, each of these species, which together represent multiple families from every major insect order, utilizes this ability as a means of navigating the environment, using learned cues to guide an active search for food and hosts, or to avoid noxious events. Nonetheless, we demonstrate not only that sedentary antlions can learn, but also, more importantly, that learning provides an important fitness benefit, namely decreasing the time to pupate, a benefit not yet demonstrated in any other species. Compared to a control group in which an environmental cue was presented randomly vis-à-vis daily prey arrival, antlions given the opportunity to associate the cue with prey were able to make more efficient use of prey and pupate significantly sooner, thus shortening their long, highly vulnerable larval stage. Whereas "median survival time," the point at which half of the animals in each group had pupated, was 46 days for antlions receiving the Learning treatment, that point never was reached in antlions receiving the Random treatment, even by the end of the experiment on Day 70. In addition, we demonstrate a novel manifestation of antlions' learned response to cues predicting prey arrival, behavior that does not match the typical "learning curve" but which is well-adapted to their sedentary predation strategy. Finally, we suggest that what has long appeared to be instinctive predatory behavior is likely to be highly modified and shaped by learning.     

Sand as a medium for transmission of vibratory signals of prey in antlions Euroleon nostras (Neuroptera: Myrmeleontidae)

Abstract.  European pit-building antlions ( Euroleon nostras / Geoffroy in Fourcroy/) detect their prey by sensing the vibrations that prey generate during locomotory activity. The behavioural reactions and some of the physical properties of substrate vibrations in sand are measured to observe signal transmission through the substrate. The frequency range of the signals of four arthropod species ( Tenebrio molitor , Pyrrhocoris apterus , Formica sp. and Trachelipus rathkei ) is 0.1–4.5 kHz and acceleration values are in the range 400 μm s⁻² to 1.5 mm s⁻². Substrate particle size and the frequency of prey signals both influence the propagation properties of vibratory signals. The damping coefficient at a frequency 300 Hz varies from 0.26 to 2.61 dB cm⁻¹ and is inversely proportional to the size of the sand particle. The damping coefficient is positively correlated with the frequency of the pulses. Vibrations in finer sand are attenuated more strongly than in coarser sand and, consequently, an antlion detects its prey only at a short distance. The reaction distance is defined as the distance of the prey from the centre of the pit when the antlion begins tossing sand as a reaction to the presence of prey. The mean reaction distance is 3.3 cm in the finest sand (particle size ≤ 0.23 mm) and 12.3 cm in coarser sand (particle size 1–1.54 mm). The most convenient sands for prey detection are considered to be medium particle-sized sands.     

Effects of larval antlions Euroleon nostras (Neuroptera, Myrmeleontidae) and their pits on the escape-time of ants

Abstract. Pit-building antlions are predators with a unique predation strategy, namely using pitfall traps constructed in loose sand to catch prey. Here, prey escape-time in the field is measured by introducing ants into one of four different treatment arenas. The first treatment lacks pits and antlions, the second includes 10 antlions that did not build pits, the third comprises eight artificially constructed pits, and the fourth is a treatment of eight antlions in pits and two without pits. Their pits are of a similar size to those used in the third treatment. When antlions are present without constructing pits, they impede the dispersal of prey. The mean escape-time for one half of the released ants is twice as long with antlions present as without them. When pits are present, the time taken for one half of the released ants to escape the predator is more than 10 times as long as when pits are absent. Escape-time from artificial pits is three times that from nonpit building antlions. Pits hinder the escape of ants and therefore increase the amount of time that the prey is available for capture. In the area where the pits are occupied by antlions, escape-time is four times longer than in a treatment with similar sized artificial pits. Thus, it appears that not only a pit, but also the presence of antlions influences the capture success.     

穴蚁蛉的研究

通过多年的观察研究,详细报导粤北穴蚁蛉的生态类型、生物学特性、个体发育与饲食频率的关系;介绍穴蚁蛉幼虫——蚁狮在治疗泌尿系统结石疾病中的重大药用价值和开发、保护蚁狮的方法和途径．     

Antlions and owlflies (Neuroptera: Myrmeleontidae,Ascalaphidae) of Kyrgyzstan

The myrmeoleontoid Neuroptera fauna of Kyrgyzstan comprises 34 species-group taxa, among which 23 are recorded from the republic for the first time. Epacanthaclisis alaica Krivokhatsky, 1998 is known only from the holotype collected in the Alai Mt. Range; Kirghizoleon cubitalis Krivokhatsky et Zakharenko, 1994 is known from Kyrgyzstan and from the Hissar Mt. country in Tajikistan; and Mesonemurus vartianorum Hölzel, 1972 is distributed from the Tien Shan to Afghanistan. 26 species are common in Kyrgyzstan and are fairly evenly distributed over its territory. New synonymies are established: Macronemurus persicus (Navás, 1915) = Barreja persica Navás, 1915, = Barreja amoena Hölzel, 1972, syn. n.; Myrmecaelurus acerbus (Walker) = Myrmeleon acerbus Walker, 1853; = Myrmecaelurus varians Navás, 1913, syn. n.; = Myrmecaelurus aequans Navás, 1913, syn. n.; = Myrmecaelurus nematicus Navás, 1932, syn. n. The polymorphous structure of the widely variable Macronemurus persicus is shown, and three its color morphs are distinguished: Macronemurus persicus var. persica (Navás, 1915), var. graciosa var. n., and var. amoena (Hölzel, 1972). Data on the antlion and owlfly fauna of some other Palaearctic countries are supplemented.     

The mitochondrial genome of Gatzara jezoensis (Neuroptera: Myrmeleontidae) and phylogenetic analysis of Neuroptera

Phylogenetic relationship within Neuroptera is controversial, particularly for the various hypotheses based on both morphological and molecular evidence. In the present study, we determined the complete mitochondrial genome (mitogenome) of Gatzara jezoensis, which is the second representative of the tribe Dendroleontini. The G. jezoensis mitogenome contained the conserved set of 37 mitochondrial genes and a putative control region, with a conserved gene arrangement which was similar to that of most sequenced neuropteran mitogenomes. All transfer RNAs exhibited the canonical cloverleaf secondary structure, except for trnS^(AGN). The control region contained two conserved elements (ploy-T stretch and ATGGTTCAAYAAAATAAYYCYCTC motif) and abundant microsatellite-like elements. The phylogenetic analysis of sequenced neuropteran mitogenomes using the concatenated protein-coding genes (PCGs) and ribosomal genes recovered the monophyly of Myrmeleontidae, which revealed this dataset could generate the more robust phylogeny of Neuroptera than that of 13 PCGs dataset.     

A genome-wide phylogeny and the diversification of genus Liriomyza (Diptera: Agromyzidae) inferred from anchored phylogenomics

The genus Liriomyza Mik (Diptera: Agromyzidae) is a diverse and globally distributed group of acalyptrate flies. Phylogenetic relationships among Liriomyza species have remained incompletely investigated and have never been fully addressed using molecular data. Here, we reconstruct the phylogeny of the genus Liriomyza using various phylogenetic methods (maximum likelihood, Bayesian inference, and gene tree coalescence) on target-capture-based phylogenomic datasets (nucleotides and amino acids) obtained from anchored hybrid enrichment (AHE). We have recovered tree topologies that are nearly congruent across all data types and methods, and individual clade support is strong across all phylogenetic analyses. Moreover, defined morphological species groups and clades are well-supported in our best estimates of the molecular phylogeny. Liriomyza violivora (Spencer) is a sister group to all remaining sampled Liriomyza species, and the well-known polyphagous vegetable pests [L. huidobrensis (Blanchard), L. langei Frick, L. bryoniae. (Kaltenbach), L. trifolii (Burgess), L. sativae Blanchard, and L. brassicae (Riley)]. belong to multiple clades that are not particularly closely related on the trees. Often, closely related Liriomyza species feed on distantly related host plants. We reject the hypothesis that cophylogenetic processes between Liriomyza species and their host plants drive diversification in this genus. Instead, Liriomyza exhibits a widespread pattern of major host shifts across plant taxa. Our new phylogenetic estimate for Liriomyza species provides considerable new information on the evolution of host-use patterns in this genus. In addition, it provides a framework for further study of the morphology, ecology, and diversification of these important flies.     

锈翅蚁蛉(脉翅目:蚁蛉科)的发育生物学研究

锈翅蚁蛉为全变态类昆虫,在贵州省一年发生1代,11月下旬开始越冬,各龄幼虫均可越冬,次年3月苏醒.在室内饲养条件下,卵期13 d±0.56 d,幼虫期63.58 d±1.81d,蛹期26.35 d±1.14 d,成虫期20 d± 1.06 d.结合野外调查和室内饲养,详细描述了锈翅蚁蛉各虫态的形态特征,幼虫的筑穴、捕食、蜕皮、化蛹、羽化以及成虫的捕食、交配等行为习性.     

Learning in a sedentary insect predator: Antlions (Neuroptera: Myrmeleontidae) anticipate a long wait

Pit-building antlions, the larvae of a winged adult insect, capture food by digging funnel-shaped pits in sand and then lying in wait, buried at the vertex, for prey to fall inside. The sedentary nature of this sit-and-wait predatory behaviour and, especially, antlions’ innate ability to detect prey arrival, do not fit the typical profile of insects that possess learning capabilities. However, we show, for the first time, that learning can play an important role in this unique form of predation. In three separate experiments, individual antlions received, once per training day, either a vibrational cue presented immediately before the arrival of food or that same cue presented independently of food arrival. Signalling of food not only produced a learned anticipatory behavioural response (Experiment 1), but also conferred a fitness advantage: Associative learning enabled antlions to dig better pits (Experiments 2 and 3), extract food more efficiently (Experiments 2 and 3), and, in turn, moult sooner (Experiment 3) than antlions not receiving the associative learning treatment.     

Thermal dependence of trap building in predatory antlion larvae (Neuroptera: Myrmeleontidae)

Trap-building predators remain under strong selection from thermal microenvironments. To address how soil temperature and body size affect trap building, we conducted a laboratory experiment using larvae of the antlion Myrmeleon bore at six ecologically relevant temperatures. Larger larvae built larger traps, and warmer soil led to more and larger traps. Body mass did not alter the dependence of trap building on temperature. Our results suggest that the physiological capacity of antlion larvae, which is affected by larval size and body temperature, is the major determinant of trap building. This effect should be considered when assessing interactions between antlions and prey.     

On Variability of Two Arabian Myrmeleontoid Lacewings (Neuroptera: Myrmeleontidae,Ascalaphidae)

Entomological Review - Body-size dimorphism in the antlion Myrmeleon hyalinus hyalinus Olivier, 1811 is described; it is manifested in the existence of two size forms resulting from different...     

Antlions (Neuroptera,Myrmeleontidae) from ornithological traps on the Curonian Spit: A three-species community containing a new species

A mixed antlion community is recorded at the “Fringilla” Research Station, “Rybachii” Biological Station, Curonian Spit, Baltic Sea, Kaliningrad Province, Russia; the adults were captured using ornithological traps and the larvae were found on sand dunes around. The ratio of the larval numbers in the mixed colonies of Myrmeleon tschernovi sp. n., Myrmeleon formicarius L., and Euroleon nostras (Geoffr.) is 100: 3: 2. The new species is described, the other two are recorded in Kaliningrad Province for the first time. Morphologically, Myrmeleon tschernovi sp. n. is similar to Myrmeleon bore (Tjed.), being its neighbor in the Baltic Region and occupying its econiche. The most characteristic distinctions between these species are found in the male genitalia and in the larval head chaetotaxy and color pattern.     

The phylogeny of lance lacewings (Neuroptera: Osmylidae)

The first phylogeny of the lacewing family Osmylidae is presented here based on a total evidence analysis of DNA sequences for multiple gene loci and morphology for representatives of almost all extant genera. Our phylogeny shows a basal dichotomy in the family, with subfamilies Protosmylinae, Spilosmylinae and Gumillinae comprising one lineage, and the other lineage including Osmylinae, Porisminae, Eidoporisminae, Kempyninae and Stenosmylinae. The status of Paryphosmylus Krüger and Lysmus Navás as members of Protosmylinae is affirmed as well as the placement of Gumillinae near Protosmylinae and Spilosmylinae. Our results suggest that Porisminae, Eidoporisminae and Stenosmylinae evolved from a common ancestor, and their relationships, including likely paraphyly of Stenosmylinae, requires further assessment. Divergence time analysis revealed that the family originated during the Late Permian before the break‐up of the supercontinent Pangaea and that present generic distributions are not due to Gondwanan biogeographic events. All major subfamily‐level lineages were present by the end of the Triassic, in agreement with the rich Mesozoic‐aged fossil record for the family.     

The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia

In the holometabolous insect order Neuroptera (lacewings), the cosmopolitan Myrmeleontidae (antlions) are the most species-rich family, while the closely related Nymphidae (split-footed lacewings) are a small endemic family from the Australian-Malesian region. Both families belong to the suborder Myrmeleontiformia, within which controversial hypotheses on the interfamilial phylogenetic relationships exist. Herein, we describe the complete mitochondrial (mt) genomes of an antlion (Myrmeleon immanis Walker, 1853) and a split-footed lacewing (Nymphes myrmeleonoides Leach, 1814), representing the first mt genomes for both families. These mt genomes are relatively small (respectively composed of 15,799 and 15,713 bp) compared to other lacewing mt genomes, and comprise 37 genes (13 protein coding genes, 22 tRNA genes and two rRNA genes). The arrangement of these two mt genomes is the same as in most derived Neuroptera mt genomes previously sequenced, specifically with a translocation of trnC. The start codons of all PCGs are started by ATN, with an exception of cox1, which is ACG in the M. immanis mt genome and TCG in N. myrmeleonoides. All tRNA genes have a typical clover-leaf structure of mitochondrial tRNA, with the exception of trnS1(AGN). The secondary structures of rrnL and rrnS are similar with those proposed insects and the domain I contains nine helices rather than eight helices, which is common within Neuroptera. A phylogenetic analysis based on the mt genomic data for all Neuropterida sequenced thus far, supports the monophyly of Myrmeleontiformia and the sister relationship between Ascalaphidae and Myrmeleontidae.     

Molecular phylogeny of the green lacewings (Neuroptera: Chrysopidae)

Abstract  The first quantitative analysis of phylogenetic relationships of green lacewings (Chrysopidae) is presented based on DNA sequence data. A single nuclear and two mitochondrial genes are used in the analysis: carbomoylphosphate synthase (CPS) domain of carbamoyl-phosphate synthetase-aspartate transcarbamoylase-dihydroorotase (CAD) (i.e. rudimentary locus), large subunit ribosomal gene (16S) and cytochrome oxidase I (COI). This study represents the first use of the CAD gene to investigate phylogenetic relationships of the lacewings. DNA sequences for 33 chrysopid species from 18 genera, representing all subfamilies and tribes, were compared with outgroups sampled from families Hemerobiidae, Osmylidae and Polystoechotidae. Parsimony analyses of the combined data set recovered all of the previously established subfamilial and tribal groups as monophyletic clades (although relatively weakly supported) except Apochrysinae sensu lato . The enigmatic Nothancyla verreauxi Navás has historically been difficult to place in a subfamily group based on morphological characteristics; molecular data presented herein do not adequately resolve this problem.     

Utilisation of prey by antlion larvae (Neuroptera: Myrmeleontidae) in terms of energy and nutrients

Prey utilisation at low prey densities was determined for third instar Cueta sp., Furgella intermedia (Markl) and Palpares annulatus (Stitz) larvae in terms of wet weight, dry weight, energy and nutrients. Prey utilisation was similar to other insects on a wet weight (42-47%), dry weight (46-49%), energetic (40-58%) and nutritive basis (62-79%). Lipids (33-36%) provided energetically the highest contribution of the nutrients ingested. The quantities of water, proteins, nucleic acids, lipids and carbohydrates extracted by the antlion larvae were in proportion to their availability in their prey, the Hodotermes mossambicus larvae. The quantities of nutrients extracted by the antlion larvae at low prey densities were not significantly influenced by the differences in mandible size, antlion body weight or the trapping method (building a pit or not) of the antlion species. It is proposed that a low metabolic rate and the accumulation of fat reserves, and not the extent of prey utilisation, enable P. annulatus larvae to tolerate a 123-d starvation period in which 22.3% of their body weight is lost.