Homology of the genital sclerites of Megaloptera (Insecta: Neuropterida) and their phylogenetic relevance

The genitalia of Megaloptera are crucial for taxonomic identification and represent a significant component of characters for phylogenetic interpretation of this order. However, several complex genital structures, especially those related to segments 9 and 11 in Megaloptera, have yet to be subjected to a comprehensive survey of homology. The terminology for genital sclerites has been variously and even incorrectly used by different authors, a fact which could lead to much confusion about character evolution. In this paper, we first present a comprehensive morphological comparison of the sclerites of male and female genital segments in 23 megalopteran genera representing all major lineages of Corydalinae, Chauliodinae and Sialidae. Accordingly, we then provide new interpretations on the homology of the genital sclerites which often appear to be considerably different among Megaloptera. Based on our new and revised homology assessments, we conclude that: (i) the small to medium‐sized sclerite beneath the ectoprocts in males of Sialidae represents the fused gonocoxites 11; (ii) the male gonocoxites 11 in Corydalidae are largely reduced and are sometimes retained as a small sclerite beneath the anus; (iii) the predominant sternite‐like sclerite of the female abdominal segment 8 represents the fused gonocoxites 8; and (iv) a pair of sclerites amalgamated with the lateral arms of male gonocoxites 10 in Chauliodinae is the gonocoxites 9. Furthermore, based on our genital homology assessments, we reconstruct an intergeneric phylogeny including all genera of Megaloptera using genital characters in a parsimonious analysis to test their phylogenetic relevance. The phylogeny herein recovered is largely congruent with the results from several previous studies, thus underlying the significant phylogenetic relevance of the megalopteran genital sclerites. The present work provides new insights into the evolution of insect genitalia.     

The genus Mesadenella (Orchidaceae) in Colombia with description of two new species

Two new species of Mesadenella, M. longipetiolata and M. bicordata, are described, illustrated and placed within the key to identification of Colombian Mesadenella species. The information about distribution and ecology of the new species as well as brief taxonomic notes are provided.     

Wing Base Structural Data Support the Sister Relationship of Megaloptera and Neuroptera (Insecta: Neuropterida)

The phylogenetic status and the monophyly of the holometabolous insect order Megaloptera has been an often disputed and long unresolved problem. The present study attempts to infer phylogenetic relationships among three orders, Megaloptera, Neuroptera, and Raphidioptera, within the superorder Neuropterida, based on wing base structure. Cladistic analyses were carried out based on morphological data from both the fore- and hindwing base. A sister relationship between Megaloptera and Neuroptera was recovered, and the monophyly of Megaloptera was corroborated. The division of the order Megaloptera, the traditional higher classification, into Corydalidae (Corydalinae + Chauliodinae) and Sialidae, was also supported by our wing base data analyses.     

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.     

Revision of the Protohermes changningensis species group from China (Megaloptera: Corydalidae: Corydalinae)

A new species group, the Protohermes changningensis group, of the dobsonfly genus Protohermes is established. Three species belonging to the new species group from China are described and illustrated, including one new species. Phylogenetic relationships among the species in this group and biogeography are discussed on the basis of a cladistic analysis using Hennig 86.     

The Neotropical genera of Acanthocinini Blanchard, 1845 (Coleoptera: Cerambycidae: Lamiinae) with the third antennomere modified

The genera of Acanthocinini which have males with antennomere III modified are discussed and redefined, a key to identify them is provided. Additionally, Alcathousites Gilmour, 1962 is proposed as a junior synonym of Xenocona Gilmour, 1960, and the following five new combinations are proposed: Xenocona asperipennis (Fairmaire & Germain, 1859) n. comb.; X. forsteri (Tippmann, 1960) n. comb.; X. penicillata (Monné, 1990) n. comb.; X. senticosa (Monné & Martins, 1976) n. comb.; and X. superstes (Erichson, 1847) n. comb. Moreover, a new species from Colombia (Huila) is described and illustrated, Xenocona antonkozlovi n. sp., and the male of X. penicillata is illustrated for the first time.     

Diving beetles of the genus Liodessus Guignot, 1953 in Colombia,with description of three new species (Coleoptera: Dytiscidae)

The Colombian species of the diving beetle genus Liodessus Guignot, 1939 Guignot, F. (1939), ‘Contribution à l’étude des Bidessus’, Bulletin de la Société d’Étude des Sciences Naturelles de Vaucluse, 10(4), 51–61. [Google Scholar] are revised. Liodessus bogotensis Guignot, 1953 Guignot, F. (1953), ‘Trente-neuvième note sur les hydrocanthares’, Revue Française d’Entomologie, 20, 109–117. [Google Scholar] is re-described. Three higher altitude species are new to science: Liodessus azufralis sp. n., L. quillacinga sp. n. and L. quimbaya sp. n. We also introduce two new subspecies, L. quillacinga cochaensis ssp. n. and L. quillacinga cumbalis ssp. n. We delineate the species using morphological structures such as male genital structure and beetle size, shape and colour pattern. Mitochondrial cox1 sequence data provided an additional character source. All the new species occur on higher altitudes above 2700 m and were collected in shallow, exposed peatland pools and puddles, mostly in Páramo. Liodessus obscurellus (LeConte, 1852), not yet recorded from Colombia, is included into the key due to its presence in nearby Costa Rica and Ecuador. The known distribution and habitat preferences of each species are outlined briefly.     

Notes on the genus Neochauliodes Weele (Megaloptera: Corydalidae) from Henan, China

The species of the genus Neochauliodes from Henan are reviewed. Three species are described as new to science: Neochauliodes digitiformis sp. nov., Neochauliodes parasparsus sp. nov. and Neochauliodes sparsus sp. nov. A key to the species from Henan is presented.     

An updated phylogeny of Deprea (Solanaceae) with a new species from Colombia: interspecific relationships,conservation assessment,and a key for Colombian species

Deprea is a neotropical genus that comprises 51 species. Recent work based on molecular data has explored its monophyly and interspecific relationships, but the relationships amongst Deprea species were not entirely elucidated. The inclusion of more accessions and molecular markers in phylogenetic analysis is likely to produce more supported hypotheses. Therefore, the main objective of this study was to perform a combined phylogenetic analysis of Deprea including seven new samples representing six species and one more DNA plastid marker than in previous studies. In that process, a new species of Deprea from Colombia was discovered and is described and illustrated here. Deprea teresitae Deanna & A. Orejuela, sp. nov. is morphologically similar to D. dilloniana and phylogenetically closely related to D. hawkesii and D. harlingiana. This new species is distinguished by the combination of short calyx lobes, non-mucronate anthers, long staminal filaments and corolla lobes, and elongated appressed fruiting calyx around a subglobose green berry. The new phylogenetic results are presented, including discussion on phylogenetic interspecific relationships, taxonomy, geographic distribution, and conservation status from D. teresitae. The synonymy of Deprea sylvarum subsp. novogranatensis under D. glabra is also proposed and an identification key to the 10 Deprea species distributed in Colombia is included.     

Review of subtribe Pterellina Rohdendorf, 1967 (Diptera: Sarcophagidae) of Middle East with descriptions of twelve new species

Summary

Based on a revision of large recent collections housed at the Natural History Museum (London), Tel Aviv University and in the private collection of Yu. Verves and L. Khrokalo (Kyiv, Ukraine), new distributional data and an updated list with comments of 28 species of Middle Eastern Sarcophagidae of the subtribe Pterellina are presented. Seventeen of them are recorded for Israel, 11 for Egypt, four for Iran, two each for the Palestinian Authority, Saudi Arabia and Turkey, and one each for Jordan and the UAE. No data are known for the territories of Bahrain, Cyprus, Iraq, Kuwait, Lebanon, Oman, Qatar, Syria and Yemen. Twelve species new to science are described, including nine from Israel (Chaetapodacra montana n. sp., Eremasiomyia bicolor n. sp., E. deserta n. sp., E. fuscata n. sp., Protomiltogramma bidentata n. sp., P. serrata n. sp., Pterella hermonica n. sp., P. indigens n. sp., P. zefatica n. sp.), two from Egypt (Chaetapodacra griseola n. sp., Eremasiomyia alba n. sp.) and one from Saudi Arabia (Protomiltogramma arabica n. sp.). The male of Eremasiomyia nigra Rohdendorf, 1935 is described for the first time. One new combination of specific name is proposed ? Protomiltogramma immunita (Villeneuve, 1923), n. comb. Original keys for the identification of genera and species are proposed. An annotated list of all world genera and species and compiled data on larval habits of Pterellina are included.     

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.     

The Protohermes fruhstorferi species group (Megaloptera: Corydalidae: Corydalinae), with description of a new species from Vietnam

A new species group, the Protohermes fruhstorferi group of the dobsonfly genus Protohermes, is recognised. Three species belonging to the new species group are described and illustrated, including one new species from northern Vietnam.     

Trichoptera fauna from Nakhon Si Thammarat Range (southern Thailand), with the description of a new species of Rhyacophila Pictet, 1834 (Trichoptera: Rhyacophilidae)

Seventy-five species of caddisflies were found near eight waterfalls in the Tai Rom Yen National Park and the Khao Luang National Park, within Nakhon Si Thammarat Range, in southern Thailand. Rhyacophila suratthaniensis sp. n. is described and figured, based on adult males. Rhyacophila suratthaniensis sp. n. is distinguished from other species of the genus by the shape of its paramere: their basal part is broader than the apical one and is separated from it by a step; apex of the paramere bears many strong spines, which are arranged like in a toothbrush. Occurrence of Trichoptera species in the observed and neighbouring areas is discussed.

http://zoobank.org/urn:lsid:zoobank.org:pub:336401B5-DAAA-4818-90EA-5DCB1944D1F0     

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.     

The adults of the Australian Megaloptera 1

A key to the supraspecific taxa of the Australian Megaloptera, as recognized in this paper, is presented. Identification of species and subspecies is possible by comparison with illustrations of genitalia which are presented for all known forms. Their geographical ranges are quoted. Species status is proposed for phaeoscius Riek and polypastus Riek, both originally described as subspecies of Archichauliodes guttiferus (Walker). A new subgenus of Archichauliodes, Apochauliodes is erected. Stenosialis Tillyard is re‐instated. The adults of the following new taxa are described: Archichauliodes cervulus, A. collifer, A. conversus, A. cuspidatus, A. isolatus, A. lewis, A. neoguttiferus. A. pictus, A. piscator, A. rieki, A. simpsoni, A. uncinatus, Protochauliodes biconicus incertus, P. b. tooloomensis, P. kirramae, Austrosialis maxmouldsi, Stenosialis hollowayi. Also presented is the hitherto undescribed female of Archichauliodes phaeoscius Riek. Austrochauliodes Riek is regarded a junior synonym of Protochauliodes Weele. Archichauliodes australica Kimmins is synonymized with Archichauliodes anagaurus Riek. Five species groups are defined and proposed for the subdivision of Archichauliodes s. str.     

The Micro-caddisflies of Lord Howe Island (Hydroptilidae: Trichoptera: Insecta)

Hydroptilid species are described from Lord Howe Island for the first time. All four species are in the endemic Australian genus Orphninotrichia, and all are endemic to Lord Howe Island. Modifications exhibited by these new species to the normally conservative Orphninotrichia features are suggestive of sympatric speciation and, for several, of a shift in larval niche.     

Contribution to the Knowledge of the Pre-imaginal Stages of Neotropical Hydrobiosidae (Trichoptera): Metachorema griseum Schmid

Mature larvae and pupae of Metachorema griseum Schmid are described for the first time. The known distribution of the genus includes the province of Neuquén in Argentina and central and southern Chile. The distinctive characters of the larvae of this genus are: (1) prothoracic sternite consisting of a wide subrectangular central sclerite and a pair of elongated lateral sclerites, (2) anterior femora with a narrow basodistal process, (3) anterior tibia and tarsus fused and (4) anal prolegs with a short lateral sclerite with basal spine and claws simple with ventral spine and two ventral setae.     

Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia

The 125 species of the satyrid butterfly tribe Pronophilini known from the three Andean Cordilleras of Colombia are listed, together with two species whose Colombian status is doubtful. The biological, ecological and biogeographical features of the tribe are described, using the listed species as examples. The features include restriction to moist conditions at high altitudes, endemism in the Cordilleras and in areas and on slopes within them, allopatric speciation and subspeciation, parapatric distributions of related species up the altitude gradients, intraspecific variation and polymorphism and apparent mimicry. The criteria used for classifying the taxa and assessing their relationships are discussed. Seventeen new species, three new subspecies and one new form are described. At the species level, 11 new synonymies are established, two synonymies are rejected, the status of 15 other taxa is revised, and 23 new combinations are made. At the generic level, two new synonymies are established, and one genus is resurrected.     

Systematics of thrips (Insecta: Thysanoptera) associated with mosses

The systematic relationships between phlaeothripine Thysanoptera species suggest that moss-feeding is an uncommon and highly derived habit. It is restricted to a single sub-tribe, the Williamsiellina, which includes two genera, Lissothrips with 17 species and Williamsiella with 25 species. Although widely distributed, these genera are particularly common in the New World. Structural variation within and between populations of these small wingless insects is discussed and the conclusion reached that species cannot be defined satisfactorily in the common North American L. muscorum -group without experimental rearing. Trisclerothrips is synonymized with Lissothrips , and Prolissothrips, Microlissothrips and Phthirothrips are synonymized with Williamsiella . One specific synonym is established, and four new species are described: L. clayae from Trinidad, L. okajimai from Japan, W. tambopata from Peru and W. zaps from Fiji.