Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

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.     

The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta)

External and internal head structures of larvae of Nevrorthidae were described in detail. The results were compared to conditions found in other representatives of Neuroptera and the other two neuropterid orders. The cladistic analysis supported the monophyly of Neuroptera, Neuroptera exclusive of Nevrorthidae, Hemerobiiformia, and Myrmeleontiformia. Neuroptera exclusive of Nevrorthidae are supported by the formation of an undivided postmentum and the presence of cryptonephric Malpighian tubules. The highly specialized articulation of the neck (Rollengelenk) and the absence of a salivary duct are autapomorphies of Nevrorthidae. Ithonidae and Polystoechotidae form a clade and are the sister group of the remaining Hemerobiiformia, which are characterized by the complete lack of a gula and a terminal filament of the antenna. Within this lineage, a clade comprising Mantispidae, Dilaridae, Berothidae, and Rhachiberothidae is well supported. Larvae of Myrmeleontiformia are characterized by a complex transformation of head structures, with a hypostomal bridge, a small triangular gula, largely reduced maxillary grooves, and anteriorly shifted posterior tentorial grooves. The slender finger‐like mid‐dorsal apodeme is another autapomorphy of the group. Psychopsidae are placed as the sister group of the remaining Myrmeleontiformia, which are characterized by a conspicuous, protruding ocular region (often less distinct or even absent in Nemopteridae). Ascalaphidae are the sister group of Myrmeleontidae. Larvae of both families share the fusion of the tibia and tarsus in the hind leg. The larval characters analysed were not sufficient for full resolution of the myrmeleontiform and hemerobiiform lineages. The position of several families such as Osmylidae, Sisyridae, and Coniopterygidae remains uncertain. The results are in agreement with an aquatic ancestor of Neuroptera and secondarily acquired terrestrial habits within the lineage (Neuroptera exclusive of Nevrorthidae), and another invasion of the aquatic environment by Sisyridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 533–562.     

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.     

Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera)

A phylogenetic analysis of Neuroptera using thirty‐six predominantly morphological characters of adults and larvae is presented. This is the first computerized cladistic analysis at the ordinal level. It included nineteen species representing seventeen families of Neuroptera, three species representing two families (Sialidae and both subfamilies of Corydalidae) of Megaloptera, two species representing two families of Raphidioptera and as prime outgroup one species of a family of Coleoptera. Ten equally most parsimonious cladograms were found, of which one is selected and presented in detail. The results are discussed in light of recent results from mental phylogenetic cladograms. The suborders Nevrorthi‐ formia, Myrmeleontiformia and Hemerobiiformia received strong support, however Nevrorthiformia formed the adelphotaxon of Myrmeleontiformia + Hemerobiiformia (former sister group of Myrmeleontiformia only). In Myrmeleontiformia, the sister‐group relationships between Psychopsidae + Nemopteridae and Nymphidae + (Myrmeleontidae + Ascalaphidae) are corroborated. In Hemerobiiformia, Ithonidae + Polystoechotidae is confirmed as the sister group of the remaining families. Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)), which has already been proposed, is confirmed. Chrysopidae + Osmylidae emerged as the sister group of a clade comprising Hemerobiidae + ((Coniopterygidae + Sisyridae) + (dilarid clade)). Despite the sister‐group relationship of Coniopterygidae + Sisyridae being only weakly supported, the position of Coniopterygidae within the higher Hemerobiiformia is corroborated. At the ordinal level, the analysis provided clear support for the hypothesis that Megaloptera + Neuroptera are sister groups, which upsets the conventional Megaloptera + Raphidioptera hypothesis.     

Phylogeny of the Neuropterida (Insecta: Holometabola)

The Neuropterida, with about 6500 known species — living fossils in a way — at the base of the Holometabola (as a sister group of the Coleoptera), comprise Raphidioptera (about 210 species, two families), Megaloptera (about 300 species, two families) and Neuroptera (6000 species, 17 families). Megaloptera + Neuroptera is argued vs. the traditional Raphidioptera + Megaloptera. Raphidioptera are undisputedly monophyletic. Monophyly of Megaloptera is the operational hypothesis, although occasionally questioned. Sucking tubes of the larvae are the most spectacular autapomorphy of Neuroptera. The construction of larval head capsules indicates three evolutionary lines: Nevrorthiformia, and Myrmeleontiformia + Hemerobiiformia. Traditional Myrmeleontiformia is Psychopsidae + (Nemopteridae + (Nymphidae + (Myrmeleontidae + Ascalaphidae))), the present approach is (Psychopsidae + Nemopteridae) + all other Myrmeleontiformia. Hemerobiiformia are based on the ‘maxillary head’ concept. The ithonid clade Ithonidae/Rapismatidae + Polystoechothidae and the dilarid clade Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)) are based on robust criteria. Other relationships remain unclear: Hemerobiidae + Chrysopidae (on similarity) and the ‘early offshoot’ concept of coniopterygidae (on autapomorphies) should not be perpetuated. Chysopidae + Osmylidae and (Hemerobiidae + (Coniopterygidae + Sisyridae)) + dilarid clade are discussed. Aquatic larvae, regarded as independent apomorphies of megaloptera and neuropteran Nevrorthidae and Sisyridae for a long time, are re‐interpreted as a synapomorphy of Megaloptera + Neuroptera and thus plesiomorphic within these groups. Terrestrial larvae (with cryptonephry to solve osmotic problems) are consequently apomorphic. Aquatic Sisyridae with cryptonephry of a single malpighian tubule, is conflicting, but larvae may have become secondarily aquatic, after a terrestrial intermezzo.     

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.     

Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola)

Abstract Segment 9 of male Raphidioptera, comprising tergite, sternite, gonocoxites, gonostyli and gonapophyses, is a benchmark for homologies in the male and female terminalia of the three Neuropterida orders Raphidioptera, Megaloptera and Neuroptera. The segments relating to genitalia are 9, 10 and 11 in males and 7, 8 and 9 in females. Results from holomorphological and recent molecular cladistic analyses of Neuropterida agree in supporting the sister‐group relationships between: (1) the Raphidioptera and the clade Megaloptera + Neuroptera, and (2) the suborder Nevrorthiformia and all other Neuroptera. The main discrepancy between the results of these studies is the nonmonophyly of the suborder Hemerobiiformia in the molecular analysis. The monophyly of the Megaloptera (which has been repeatedly questioned) is further corroborated by a hitherto overlooked ground pattern autapomorphy: the presence of eversible sacs within the complex of the fused gonocoxites 11 in Corydalidae and Sialidae. The recently discovered paired complex of gonocoxites 10 (parameres) in Nipponeurorthus (Nevrorthidae) indicates that the curious apex of sternite 9 of Nevrorthus and Austroneurorthus is the amalgamation of the sclerites of gonocoxites 10 with sternite 9, interpreted as synapomorphic. In the molecular study, the Nevrorthidae, Sisyridae and Osmylidae branch off in consecutive splitting events, a result that is supported by the analysis of male genital sclerites reported here. Extraordinary parallel apomorphies (e.g. excessive enlargement and modification of gonocoxites 10 ending in a thread‐like ‘penisfilum’) in derived representatives of Coniopterygidae, Berothidae, Rhachiberothidae and Mantispidae corroborate the dilarid clade of the morphological analysis and leads us to hypothesize a sister‐group relationship of the Coniopterygidae with the dilarid clade. A re‐interpretation of the tignum of Chrysopidae as gonocoxites 11 means that the structure previously called the gonarcus represents the fused gonocoxites 9. In Hemerobiidae, the corresponding sclerite is consequently also homologized as fused gonocoxites 9. The enlargement of the lateral wings of the gonocoxites in both families is interpreted as a synapomorphy. Excessive enlargement of gonostyli 11 in the Polystoechotid clade and Myrmeleontiformia supports a sister‐group relationship of these two clades. The occurrence of certain serial homologues of female genitalia structures (gonocoxites and gonapophyses), such as the digitiform processus together with the flat appendices in segment 8 of certain Myrmeleontidae, or the wart‐like processus together with the flat circular sclerites in segment 7 of certain Berothidae, as well as the presence of gonocoxites 8 as pseudosternites in certain Nemopteridae and Coniopterygidae, are probably character reversals. The digitiform processus of tergite 9 (pseudogonocoxites) in Rhachiberothidae and Austroberothella (Berothidae) are either independently developed acquisitions with a function in oviposition, or are homologous sclerites, possibly of epipleurite origin.     

Seasonality in adult flight activity of two neuroptera assemblages of southern Mali

The seasonality of insect assemblages in Africa is poorly investigated. To provide information on the relationships between climate and insect assemblages in the Sudanian region, strongly affected by climate change, we studied Myrmeleontidae and Ascalaphidae assemblages (Insecta: Neuroptera) for 7 and 5 consecutive years respectively in southern Mali. To make the species inventory as exhaustive as possible, we performed weekly sampling by netting and light trapping. For both assemblages, results showed very similar patterns of variation in species diversity throughout the year. Adults of Myrmeleontidae and Ascalaphidae were active all year, and the species succession was influenced by a strong temporal segregation. Species diversity peaked at the end of the rainy season and surprisingly during the dry season. Principal component analysis of the climatic factors followed by co‐inertia analysis applied to two data sets, one comprising climatic factors and the other reporting presence/absence of species, showed a good association between the annual trend of climatic factors and the species diversity. But no well defined species grouping was clearly linked to a particular period of the year. This tight association between climate and species composition suggests that even small climate changes could modify significantly species assemblage characteristics.     

A new Upper Carboniferous stem‐orthopteran (Insecta) from Ningxia (China)

Abstract During the Upper Carboniferous, orthopteran insects (grasshoppers, katydids, and crickets) were represented by numerous species distantly related to crown‐orthopterans, such as lobeattid and cnemidolestodean insects. The panorthopterans, including total‐orthopterans and their closest relatives, are represented by comparatively rarer species in localities of this period. Here we describe Heterologus duyiwuer sp. nov., an infrequent panorthopteran from the Late Carboniferous locality of Xiaheyan Village (Zhongwei City, Ningxia Hui Autonomous Region, China). The only available specimen is composed of an isolated forewing exhibiting a combination of character states previously unknown, in particular the lack of posterior radius (RP) / anterior Media (MA) connection, late branchings of the media (M) and anterior cubitus (CuA), and a branched posterior branch of the posterior cubitus (CuPb). Based on its unusual branching pattern, the composite stem resulting from the fusion of CuA and CuPaα (second anterior branch of CuP) is assumed to be composed of a branched CuA and a simple CuPaα.     

Owlflies are derived antlions: anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera)

The first phylogenomic analysis of the antlions is presented, based on 325 genes captured using anchored hybrid enrichment. A concatenated matrix including 207 species of Myrmeleontoidea (170 Myrmeleontidae) was analysed under maximum likelihood and Bayesian inference. Both Myrmeleontidae (antlions) and Ascalaphidae (owlflies) were recovered as paraphyletic with respect to each other. The majority of the subfamilies traditionally assigned to both Myrmeleontidae and Ascalaphidae were also recovered as paraphyletic. By contrast, all traditional antlion tribes were recovered as monophyletic (except Brachynemurini), but most subtribes were found to be paraphyletic. When compared with the traditional classification of Myrmeleontidae, our results do not support the current taxonomy. Therefore, based on our phylogenomic results, we propose a new classification for the antlions, which synonymizes Ascalaphidae with Myrmeleontidae and divides the family into four subfamilies (Ascalaphinae, Myrmeleontinae, Dendroleontinae and Nemoleontinae) and 17 tribes. We also highlight the most pressing issues in antlion systematics and indicate taxa that need further taxonomic and phylogenetic attention. Finally, we present a comprehensive table placing all extant genera of antlions and owlflies in our new proposed classification, including details on the number of species, distribution and notes on the likely monophyly of each genus.     

Traits and evolution of wing venation pattern in paraneopteran insects

Two different patterns of wing venation are currently supposed to be present in each of the three orders of Paraneoptera. This is unlikely compared with the situation in other insects where only one pattern exists per order. We propose for all Paraneoptera a new and unique interpretation of wing venation pattern, assuming that the convex cubitus anterior gets fused with the common stem of median and radial veins at or very near to wing base, after separation from concave cubitus posterior, and re‐emerges more distally from R + M stem. Thereafter, the vein between concave cubitus posterior and CuA is a specialized crossvein called “cua‐cup,” proximally concave and distally convex. We show that despite some variations, that is, cua‐cup can vary from absent to hypertrophic; CuA can re‐emerge together with M or not, or even completely disappear, this new interpretation explains all situations among all fossil and recent paraneopteran lineages. We propose that the characters “CuA fused in a common stem with R and M”and “presence of specialized crossvein cua‐cup” are venation apomorphies that support the monophyly of the Paraneoptera. In the light of these characters, we reinterpret several Palaeozoic and early Mesozoic fossils that were ascribed to Paraneoptera, and confirm the attribution of several to this superorder as well as possible attribution of Zygopsocidae (Zygopsocus permianus Tillyard, 1935) as oldest Psocodea. We discuss the situation in extinct Hypoperlida and Miomoptera, suggesting that both orders could well be polyphyletic, with taxa related to Archaeorthoptera, Paraneoptera, or even Holometabola. The Carboniferous Protoprosbolidae is resurrected and retransferred into the Paraneoptera. The genus Lithoscytina is restored. The miomopteran Eodelopterum priscum Schmidt, 1962 is newly revised and considered as a fern pinnule. In addition, the new paraneopteran Bruayaphis oudardi gen. nov. et sp. nov. is described fromthe Upper Carboniferous of France (see Supporting Information). J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

NEW FOSSIL LACEWINGS AND ANTLIONS (INSECTA, NEUROPTERA) FROM THE LOWER CRETACEOUS CRATO FORMATION OF BRAZIL

Abstract:  Remarkable new fossil taxa of Neuroptera from the laminated limestone of the Crato Formation, north-east Brazil, are described: Nuddsia longiantennata gen. et sp. nov. (Osmylidae, Gumillinae), the first fossil record of this family from South America, Parapalaeoleon magnus gen. et sp. nov. (Palaeoleontidae), and Triangulochrysopa formosa sp. nov. (Mesochrysopidae). A diagnosis of Gumillinae is provided; Epiosmylidae is considered to be a synonym of this subfamily. A revised diagnosis of Palaeoleontidae is provided, based on a new interpretation of venational characters evident on the exceptionally well-preserved specimen of P. magnus ; vein homologies are determined and the fusion of MP and CuA is discussed. The genus Triangulochrysopa has been known previously only from the Lower Cretaceous of Las Hoyas, Spain.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Species richness and composition of Neuroptera in the forests fregments of the Taurus Mountains Range,Turkey

In this study, diversity, species richness and composition of Neuroptera has been studied in the forest edges and fragments in the Taurus Mountain Range, southern Turkey. Sampling for species collection was carried out from April 2017 to September 2018 at different distances from the forest center, i.e., (0–500 m), forest mid-interior (501–1000 m), and forest edge (1001–3000 m). A total of 975 adults were collected frequently belonging to the families Ascalaphidae, Coniopterygidae, and Hemerobiidae from the forest edges while Chrysopidae and Myrmeleontidae were most common along the mid-interior regions of the forest. Majority of adutls caught from the mid-interior region comprised of female adults while the males of most species were abundant along the forest edges. Although the forest center shows the largest value for the Dominance species-diversity index and the smallest value for the Shannon index, forest edge was found highest for the Simpson index. The abundance of Neuroptera decreased with wind speed but increased with the temperature in the edge regions. Principal Component Analysis (PCA) indicated that some environmental and habitat variables, e.g. wind speed, temperature, and distance to the forest center, mid-interior and edge, accounted for species distribution patterns in Neuroptera. In the forest center, a linear correlation between wind speed, temperature and specimen abundance was recorded, while these factors were found negatively correlated with specimens abundance in the mid-interior regions of the forest.     

Phylogeny of the Neuropterida: a first molecular approach

Abstract. In a first molecular approach specially dedicated to examining the phylogeny of the Neuropterida, two nuclear and two mitochondrial genes were tested: 18S rRNA, translation elongation factor‐1α, cytochrome c oxidase subunit 3 and 16S rRNA. Molecular results are discussed in the light of a previous holomorphological cladistic analysis. The hypothesis of a sister‐group relationship Raphidioptera + (Neuroptera + Megaloptera) put forward in recent morphological analyses is supported by our data, which is in contrast to the traditional view (Raphidioptera + Megaloptera) + Neuroptera. Furthermore, the Nevrorthidae (constituting the suborder Nevrorthiformia) as a sister group of all other Neuroptera is confirmed. The disruption of the suborder Hemerobiiformia is the most conflicting result of the molecular analysis. Sisyridae and Osmylidae do not cluster within Hemerobiiformia, but represent two distinct and widely separated branches. The remaining Hemerobiiformia emerge as the sister group of the suborder Myrmeleontiformia, which is once more confirmed as monophyletic. Among the genes tested, cytochrome c oxidase subunit 3 proved to be most potent for resolving the phylogenetic relationships among Neuropterida. The nuclear gene for the ribosomal 18S rRNA is too conserved within the alignable regions, whereas the variable sections are too divergent to be applicable within this evolutionary time frame. The elongation factor‐1α gene proved to exist in more than one copy in Neuropterida, and thus is not applicable in the present state of knowledge. With respect to the mitochondrial sequences (cytochrome c oxidase subunit 3, 16S rRNA), saturation impedes the unambiguous resolution of deeper nodes. Apparently, due to early diversification of the heterogeneous Neuroptera, phylogenetic analysis of this group remains a challenge with respect to selection of the proper genes and mutatis mutandis the morphological approach.     

Eversible abdominal vesicles and some observations of the male reproductive system of the spoon wing lacewing Palmipenna (Neuroptera: Nemopteridae)

The anatomy and histology of the abdominal eversible vesicles and the male reproductive tract of the spoonwing lacewing Palmipenna (Neuroptera: Nemopteridae) have been examined. The eversible vesicles open as a pair of large bulbous sacs between tergites five and six, each folding into halves during retraction. They consist of highly pleated cuticle, beneath which are typical gland cells, each having a circular or oval end apparatus surrounded by closely packed microvilli. These communicate to the surface via cuticularized channels. In spite of considerable behavioral observations, male Palmipenna were never noted with everted vesicles. Even during mating trials, where females were presented to males in the field, the vesicles were never everted during the attempted copulation that ensued. Our observations indicate that mate attraction is mediated by the release of a female pheromone. The function of the eversible vesicles and their associated gland cells remains unknown, and their structure appears to be unique to the Nemopteridae. The reproductive tract is similar to that of other Neuroptera, consisting of a pair of five-lobed testes, a medium-to-large pair of seminal vesicles, and three pairs of accessory glands. The major accessory glands are surrounded by circular and longitudinal muscle, and are lined by an epithelium, the cells of which presumably secrete the amorphous rods of material always present in this pair of glands. The sperm in the seminal vesicles are elongate, with a pointed head and a 9 + 9 + 2 configuration in the flagellum. A single spermatophore, similar in shape to that described for other Neuroptera, was found occluding the bursa copulatrix of a teneral female. © 1994 Wiley-Liss, Inc.     

Discovery of the genus Suhpalacsa Lefèbvre (Neuroptera: Ascalaphidae: Ascalaphinae) in Japan, with description of a new species

Suhpalacsa iriomotensis sp. nov. (Neuroptera: Ascalaphidae: Ascalaphinae), is described from Iriomotejima Island, Japan. This new species can be easily distinguished from other species of the genus by the unique ventrolateral prominences of the male ectoproct. This is the first record of the genus Suhpalacsa from Japan. The morphology of the male ectoproct and the monophyly of Suhpalacsa are discussed briefly.     

Subtle variation in size and shape of the whole forewing and the red band among co‐mimics revealed by geometric morphometric analysis in Heliconius butterflies

Heliconius are unpalatable butterflies that exhibit remarkable intra‐ and interspecific variation in wing color pattern, specifically warning coloration. Species that have converged on the same pattern are often clustered in Müllerian mimicry rings. Overall, wing color patterns are nearly identical among co‐mimics. However, fine‐scale differences exist, indicating that factors in addition to natural selection may underlie wing phenotype. Here, we investigate differences in shape and size of the forewing and the red band in the Heliconius postman mimicry ring (H. erato phyllis and the co‐mimics H. besckei, H. melpomene burchelli, and H. melpomene nanna) using a landmark‐based approach. If phenotypic evolution is driven entirely by predation pressure, we expect nonsignificant differences among co‐mimics in terms of wing shape. Also, a reinforcement of wing pattern (i.e., greater similarity) could occur when co‐mimics are in sympatry. We also examined variation in the red forewing band because this trait is critical for both mimicry and sexual communication. Morphometric results revealed significant but small differences among species, particularly in the shape of the forewing of co‐mimics. Although we did not observe greater similarity when co‐mimics were in sympatry, nearly identical patterns provided evidence of convergence for mimicry. In contrast, mimetic pairs could be distinguished based on the shape (but not the size) of the red band, suggesting an “advergence” process. In addition, sexual dimorphism in the red band shape (but not size) was found for all lineages. Thus, we infer that natural selection due to predation by birds might not be the only mechanism responsible for variation in color patterns, and sexual selection could be an important driver of wing phenotypic evolution in this mimicry ring.