Comparing head muscles among Drusinae clades (Insecta: Trichoptera) reveals high congruence despite strong contrasts in head shape.

The subfamily Drusinae (Limnephilidae, Trichoptera) comprises a range of species exhibiting differently shaped head capsules in their larval stages. These correspond to evolutionary lineages pursuing different larval feeding ecologies, each of which uses a different hydraulic niche: scraping grazers and omnivorous shredders sharing rounded head capsules and filtering carnivores with indented and corrugated head capsules. In this study, we assess whether changes in head capsule morphology are reflected by changes in internal anatomy of Drusinae heads. To this end, internal and external head morphology was visualized using µCT methods and histological sections in three Drusinae species-Drusus franzi, D. discolor and D. bosnicus-representing the three evolutionary lineages. Our results indicate that Drusinae head musculature is highly conserved across the evolutionary lineages with only minute changes between taxa. Conversely, the tentorium is reduced in D. discolor, the species with the most aberrant head capsule investigated here. Integrating previous research on Drusinae head anatomy, we propose a fundamental Drusinae blueprint comprising 29 cephalic muscles and discuss significance of larval head capsule corrugation in Trichoptera.

Morphological description and DNA-based association of the last instar larva of Erotesis schachti Malicky 1982 (Trichoptera: Leptoceridae), an endemic of the Iberian Peninsula.

In this paper we describe the main morphological characteristics that distinguish the full-grown larva of Erotesis schachti, an endemic of the Iberian Peninsula. The conspecificity of the larva and adult was confirmed by DNA analysis. Morphological features that easily discriminate it from the similar species Erotesis baltica are given.

On the value of Burmese amber for understanding insect evolution: Insights from †Heterobathmilla - an exceptional stem group genus of Strepsiptera (Insecta).

Burmese amber and amber from other periods and regions became a rich source of new extinct insect species and yielded important insights in insect evolution in the dimension of time. Amber fossils have contributed to the understanding of the phylogeny, biology, and biogeography of insects and other groups, and have also gained great importance for dating molecular trees. Another major potential is the documentation of faunal, floral and climatic shifts. Evolutionary transitions can be well-documented in amber fossils and can reveal anatomical transformations and the age of appearance of structural features. Here, using a new stem group species of Strepsiptera from Burmite, we evaluate this potential of amber insect fossils to assess the current phylogeny of Strepsiptera, with the main emphasis on the early splitting events in the stem group. Amber fossils have greatly contributed to the understanding of the evolution of Strepsiptera in the late Mesozoic and the Cenozoic. †Heterobathmilla kakopoios Pohl and Beutel gen. et sp. n. described here is placed in the stem group of the order, in a clade with †Kinzelbachilla (†Kinzelbachillidae) and †Phthanoxenos (†Phthanoxenidae). †Phthanoxenidae has priority over †Kinzelbachillidae, and the latter is synonymised. The superb details available from this new fossil allowed us to explore unique features of the antennae, mouthparts, and male copulatory apparatus, and to provide a phylogenetic hypothesis for the order. The younger †Protoxenos from Eocene Baltic amber was confirmed as sister to all remaining extinct and extant groups of Strepsiptera, whereas the position of the Cretaceous †Cretostylops in the stem group remains ambivalent. While the value of Burmite and amber from other periods has a recognized impact on our knowledge of the evolution in various lineages, this new fossil does not fundamentally change our picture of the phylogeny and evolution of early Strepsiptera. However, it offers new insights into the morphological diversity in the early evolution of the group.

Identification and morphology of two polycentropodid caddisfly larvae from the Cevennes and the French and Italian Alps: Plectrocnemia cevennensis Coppa 2011 and Plectrocnemia praestans McLachlan 1884.

The collection of adults, mature male pupae, and the absence of confusing species at the sampling sites located in the Cevennes and French Alps enabled descriptions of the hitherto unknown larvae of Plectrocnemia cevennensis Coppa 2011 and P. praestans McLachlan 1884 (Trichoptera, Polycentropodidae). We present information on the morphology of the larvae and illustrate the most important diagnostic features. In the context of the larval key to European Plectrocnemia species by Waringer Graf (2020), P. cevennensis keys together with P. scruposa McLachlan 1880, whereas P. praestans keys with P. conspersa (Curtis 1834). These two species pairs are morphologically very close; P. cevennensis and P. scruposa can be separated by the numbers of secondary setae on the last abdominal sternum, but the separation of P. praestans and P. conspersa is not always possible.

The larva of Sericostoma galeatum Rambur 1842 (Trichoptera: Sericostomatidae), based on French material.

The present paper describes the previously unknown larva of Sericostoma galeatum based on French material. Information on the morphology of the larva is given, and the most important morphological characters are illustrated. Sericostoma galeatum is very close to the S. personatum / S. flavicorne species pair and cannot be separated yet from those confusing congeners. With respect to the distribution, S. galeatum is confined to the French, Italian, and Swiss western Alps and the Apennines. In addition, ecological information is provided.

Comparative morphology of extant raptorial Mantispoidea (Neuroptera: Mantispidae, Rhachiberothidae) suggests a non-monophyletic Mantispidae and a single origin of the raptorial condition within the superfamily.

Adult external morphology of the extant raptorial Mantispoidea (Insecta: Neuroptera: Mantispidae and Rhachiberothidae) is compared emphasizing the morphology of the subfamily Symphrasinae as a key group to understand the phylogenetic relationships among the members of the superfamily. Plega dactylota Rehn, 1939 is thoroughly characterized in order to exemplify the morphology of the Symphrasinae. Additionally, following a review of the literature and examination of comparative material of Dilaridae, Berothidae, Rhachiberothidae and all Mantispidae subfamilies, a new interpretation of the components of the raptorial apparatus (i.e., head, prothorax, grasping forelegs, as well as integumentary specializations) is presented. Also, wing venation for these groups is reinterpreted, and new homology hypotheses for wing venation are proposed based on tracheation and comparative analyses. Given the high morphological divergence on the genital sclerites within the Mantispoidea, plus the confusing previous usage of neutral terminology and terms referring to appendages across taxonomic and morphological studies, we attempt to standardize, simplify, and situate terminology in an evolutionary context under the "gonocoxite concept" (multi-coxopod hypothesis). The remarkable morphological similarity of the genital sclerites of Symphrasinae and Rhachiberothidae (sensu U. Aspöck Mansell 1994) with the Nallachinae (Dilaridae) was taken as a starting point to understand the morphology of other Mantispidae subfamilies. Based on these morphological comparisons, we provide a revised phylogenetic analysis of Mantispoidea. This new phylogenetic analysis supports a sister group relationship between the family Rhachiberothidae, comprising Rhachiberothinae and Symphrasinae, and the family Mantispidae, including the subfamily Mantispinae and its sister taxa Drepanicinae and Calomantispinae, which may represent a single subfamily. Based on these analyses, raptorial condition probably evolved a single time in these insects and subsequently became diversified in the two sister clades of the raptorial Mantispoidea.

Taxonomic notes on owlflies from Pakistan (Neuroptera: Myrmeleontidae: Ascalaphinae).

Taxonomic notes are presented on the former Ascalaphidae (owlflies), now subfamily Ascalaphinae of the family Myrmeleontidae from Pakistan. An updated checklist of new records is provided that accounts for all known 15 genera and 22 species from Pakistan, excluding the species of the tribe Palparini. Geographical distribution maps and an updated identification key to all known genera and species from Pakistan are also given. We synonymized two monotypic genera, Horischema Mészáros Ábrahám, 2003 and Perissoschema Mészáros Ábrahám, 2003 both as junior synonyms of Ogcogaster Westwood, 1847 based on very similar male genitalia and external morphological characters. Moreover, Perissoschema evae Mészáros Ábrahám, 2003 is treated as a junior synonym of Ogcogaster segmentator Westwood, 1847. We recorded three genera for the first time from Pakistan, i.e., Acheron Lefèbvre, 1842, Glyptobasis McLachlan, 1873, and Suphalomitus van der Weele, 1909. To more completely resolve the generic or specific status of those poorly known species from Pakistan, broader sampling throughout the country is needed.

A Revised Description of the Larva of Leptocerina pauliani (Ross 1957 (Trichoptera: Leptoceridae), an Endemic Species of La Réunion Island.

Leptocerina is an endemic Afrotropical genus of Trichoptera, with 19 species documented but only five partially described at the larval stage. The first description of Leptocerina pauliani (Ross 1957) larva by Marlier Marlier (1982) included very few illustrations, making comparisons with other species challenging. I present a revised description of the L. pauliani larval stage, using specimens collected from the River Marsouins in La Réunion Island. I apply a focus stacking approach for emphasizing coloration patterns and providing a detailed description of larval characters. Criteria for distinguishing L. pauliani from the other species of Leptocerina already described or partially described for the larval stage are discussed.

Description of three new species of Psilotreta (Trichoptera: Odontoceridae) from Japan.

Three new species of the caddisfly genus Psilotreta (Trichoptera: Odontoceridae) from Japan are described: P. voluta sp. nov., P. moritai sp. nov., and P. flavida sp. nov. These three species are easily distinguishable by the male genitalia from previously known Japanese species, P. japonica (Banks 1906) and P. kisoensis Iwata 1928. The three new species are similar to P. vertebrata Yuan et al. 2008, P. cuboides Yuan et al. 2008, and P. excavata Yuan et al. 2008 from the Chinese mainland, and P. clyssan Malicky 2014 from Taiwan, in the shapes of the lateral processes and intermediate appendages of segment X of the male genitalia, but can be distinguished from these species and each other. All Japanese species of Psilotreta listed above belong to the P. chinensis Species Group.

A new species, Agapetus kaengkrungensis (Trichoptera: Glossosomatidae) from Kaeng Krung National Park, southern Thailand with the distribution map of the genus in Thailand.

The male of a new species of caddisfly, Agapetus kaengkrungensis n. sp. (Glossosomatidae) is described and illustrated from Kaeng Krung National Park, Surat Thani Province, southern Thailand. Agapetus kaengkrungensis n. sp. is distinguished from other species by the characters of segment IX and inferior appendages. The distributions of the Agapetus spp. of Thailand are mapped and discussed.

A new species of Setodes Rambur (Trichoptera: Leptoceridae) from northern Michigan, USA.

Setodes truncatus n. sp. is illustrated and described herein based on specimens collected from 2 adjacent watersheds of the Huron Mountains in northern Michigan (USA). Both known populations inhabit undisturbed sandy rivers. Males of S. truncatus appear most similar to those of S. incertus (Walker) but can be separated based on their short and truncate inferior appendages and by other differences in the genitalia. The larva and female both remain unknown. A revised key to males of the 10 North American species of Setodes is provided.

Anatomy and histology of the alimentary canal of larvae and adults of Chrysoperla externa (Hagen, 1861) (Neuroptera: Chrysopidae).

The larvae of the lacewing Chrysoperla externa are important predators with the potential to be used in the biological control in agriculture. Although some studies provide important data on the gut morphology in lacewings, they are limited to few species. This study describes the anatomy and histology of the alimentary canal in the predatory larvae and herbivorous adult of C. externa. In larvae, the crop is the larger part of the foregut and it is connected to the midgut by the stomodeal valve. The midgut is an enlarged sac-like organ. At the mid-hindgut transition, there are eight Malpighian tubules. The hindgut is a non-functional vestigial region in the larvae. In adults, the crop has a diverticulum associated with large tracheal trunks, a conic proventriculus with sclerotized lips followed by an elongated tubular midgut. Histological analyses of larval and adult midgut show the presence of a single-layered epithelium with columnar cells with well-developed brush border, nests of regenerative cells, and a peritrophic matrix lining the midgut lumen. The hindgut in adults has an epithelium with cubic cells lined by a thin cuticular intima and rectal pads in the rectum. These data are discussed in comparison with the digestive tract in other Chrysopidae.

The endoparasitic larval stages of Eoxenos laboulbenei: An atypical holometabolan development (Strepsiptera, Mengenillidae).

Endoparasitic larval stages of Eoxenos laboulbenei were documented with different techniques, with a main focus on the male tertiary larva. Two discrete endoparasitic stages occur, the secondary and the tertiary larva. The presence of large compound eyes and externally visible wing buds in the tertiary larva is a unique feature within Holometabola. The brain with large optic lobes is followed by a single postcephalic ganglionic complex. The cephalic musculature is greatly reduced but pharyngeal dilators and muscles associated with the mouth field are present. Postcephalic sclerites are absent except for the pronotum. The segmented legs bear filiform pretarsal claws. The indirect flight muscles fill up a large part of the metathorax. The 10-segmented abdomen lacks appendages. Pleural folds are present on the thorax and abdomen. The digestive tract is characterized by a very short oesophagus. The large midgut and the narrow hindgut are disconnected. Six short Malpighian tubules are present. Large testes fill out almost the entire abdomen. In contrast to the tertiary larva, the muscles of the secondary larva are not fully differentiated. Cephalic appendages are present as bud-shaped anlagen. The legs lack a pretarsal claw. The developmental transformations are outlined and discussed, also with respect to phylogenetic implications.

Habitat divergence shapes the morphological diversity of larval insects: insights from scorpionflies.

Insects are the most diverse group of organisms in the world, but how this diversity was achieved is still a disputable and unsatisfactorily resolved issue. In this paper, we investigated the correlations of habitat preferences and morphological traits in larval Panorpidae in the phylogenetic context to unravel the driving forces underlying the evolution of morphological traits. The results show that most anatomical features are shared by monophyletic groups and are synapomorphies. However, the phenotypes of body colorations are shared by paraphyletic assemblages, implying that they are adaptive characters. The larvae of Dicerapanorpa and Cerapanorpa are epedaphic and are darkish dorsally as camouflage, and possess well-developed locomotory appendages as adaptations likely to avoid potential predators. On the contrary, the larvae of Neopanorpa are euedaphic and are pale on their trunks, with shallow furrows, reduced antennae, shortened setae, flattened compound eyes on the head capsules, and short dorsal processes on the trunk. All these characters appear to be adaptations for the larvae to inhabit the soil. We suggest that habitat divergence has driven the morphological diversity between the epedaphic and euedaphic larvae, and may be partly responsible for the divergence of major clades within the Panorpidae.

Contulma vascaina sp. n. (Insecta: Trichoptera: Anomalopsychidae), a new species from Rio de Janeiro state, southeastern Brazil.

The caddisfly genus Contulma Flint, 1969, with 30 described species, is endemic to the Neotropics. In addition to the five species currently recorded from Brazil, a new species of Contulma is described and illustrated in this work. Contulma vascaina sp. n. is similar to C. sana Jardim and Nessimian, 2011 in prominent, slender and rod-like posterolateral lobe of segment IX, but the new species has a shorter posterolateral lobe, with long, spine-like seta apically. Contulma vascaina sp. n. can also be characterized by the presence of a large, hood-like sclerotized plate on the endothecal phallic membrane. The female genital structure is also described and illustrated.

New long-proboscid lacewings of the mid-Cretaceous provide insights into ancient plant-pollinator interactions.

Many insects with long-proboscid mouthparts are among the pollinators of seed plants. Several cases of the long-proboscid pollination mode are known between fossil insects (e.g., true flies, scorpionflies, and lacewings) and various extinct gymnosperm lineages, beginning in the Early Permian and increasing during the Middle Jurassic to Early Cretaceous. However, details on the morphology of lacewing proboscides and the relevant pollination habit are largely lacking. Here we report on three lacewing species that belong to two new genera and a described genus from mid-Cretaceous (Albian-Cenomanian) amber of Myanmar. All these species possess relatively long proboscides, which are considered to be modified from maxillary and labial elements, probably functioning as a temporary siphon for feeding on nectar. Remarkably, these proboscides range from 0.4-1.0 mm in length and are attributed to the most diminutive ones among the contemporary long-proboscid insect pollinators. Further, they clearly differ from other long-proboscid lacewings which have a much longer siphon. The phylogenetic analysis indicates that these Burmese long-proboscid lacewings belong to the superfamily Psychopsoidea but cannot be placed into any known family. The present findings represent the first description of the mouthparts of long-proboscid lacewings preserved in amber and highlight the evolutionary diversification of the ancient plant-pollinator interactions.