An integrative approach to the study of Kempnyia Klapálek, 1914 (Plecoptera: Perlidae) from Brazil: Support for the description of four new species and a basis for future studies.

Kempnyia (Plecoptera: Perlidae) is an endemic genus of Brazilian stoneflies that has 36 valid species and is distributed primarily in the Atlantic Forest and the mountainous areas of Central Brazil, particularly in Goiás and Tocantins states. Despite being the Brazilian genus with the most DNA sequences available on GenBank, integrative studies on the genus began only recently, in 2014. In this context, herein we studied the morphology and molecular data of Kempnyia specimens deposited in the Aquatic Biology Laboratory (UNESP, Assis) and the Entomology Museum of the Federal University of Viçosa (UFVB, Viçosa) collections. For the integrative approach adopted, in addition to studying the specimens morphologically, we used sequences of the COI mitochondrial gene combined with the following species delimitation methods: Automatic Barcode Gap Discovery (ABGD), both primary (ABGDp) and recursive (ABGDr) partitions; Assemble Species by Automatic Partitioning (ASAP); Poisson Tree Processes (PTP) and the Bayesian implementation of the Poisson Tree Processes (bPTP). As a result, we provided 28 new COI sequences of 21 species and support the description of four new species, namely, K. guarani sp. nov., K. tupiniquim sp. nov., K. una sp. nov., and K. zwickii sp. nov., consequently increasing the known diversity of the genus to 40 species. We also discuss the morphological variations observed in other species of the genus and provide several new geographic records. Therefore, our study brings new insights into the values of intra- and interspecific molecular divergence within Kempnyia, serving as a basis for new studies.

Morphology of the pterothoracic musculature in Paraneoptera and its phylogenetic implication (Insecta: Neoptera).

Although the monophyly of Paraneoptera (=hemipteroid orders or Acercaria, composed of Psocodea, Thysanoptera and Hemiptera) has been widely accepted morphologically, the results from molecular phylogenetic and phylogenomic analyses contradict this hypothesis. In particular, phylogenomic analyses provide strong bootstrap support for the sister group relationship between Psocodea and Holometabola, that is, paraphyly of Paraneoptera. Here, we examined the pterothoracic musculature of Paraneoptera, as well as a wide range of other neopterous insect orders, and analysed its phylogenetic implication. By using the synchrotron microcomputed tomography (µCT) and parsimony-based ancestral state reconstruction, several apomorphic conditions suggesting the monophyly of Paraneoptera, such as the absence of the II/IIItpm7, IIscm3, IIIspm2 and IIIscm3 muscles, were identified. In contrast, no characters supporting Psocodea + Holometabola were recovered from the thoracic muscles. These results provide additional support for the monophyly of Paraneoptera, together with the previously detected morphological apomorphies of the head, wing base, and abdomen.

Anatomical, phenological and genetic aspects of the host-parasite relationship between Andrena vaga (Hymenoptera) and Stylops ater (Strepsiptera).

Stylops ater is an endoparasite of the mining bee Andrena vaga with extreme sexual dimorphism and hypermetamorphosis. Its population structure, parasitization mode, genetic diversity and impact on host morphology were examined in nesting sites in Germany to better understand this highly specialized host­parasite interaction. The shift in host emergence due to stylopization was proven to be especially strong in A. vaga. Around 10% of bees hosted more than 1 Stylops, with at maximum 4. A trend in Stylops' preference for hosts of their own sex and a sex-specific position of extrusion from the host abdomen was found. Invasion of Andrena eggs by Stylops primary larvae was depicted for the first time. Cephalothoraces of female Stylops were smaller in male and pluristylopized hosts, likely due to lower nutrient supply. The genes H3, 18S and cytochrome c oxidase subunit 1 were highly conserved, revealing near-absence of local variation within Stylops. Ovaries of hosts with male Stylops contained poorly developed eggs while those of hosts with female Stylops were devoid of visible eggs, which might be due to a higher protein demand of female Stylops. Male Stylops, which might have a more energy-consuming development, led to a reduction in head width of their hosts. Host masculinization was present in the leaner shape of the metabasitarsus of stylopized females and is interpreted as a by-product of manipulation of the host's endocrine system to shift its emergence. Stylopization intensified tergal hairiness, most strongly in hosts with female Stylops, near the point of parasite extrusion, hinting towards substance-induced host manipulation.

Behavioural function and development of body-to-limb proportions and active movement ranges in three stick insect species.

Overall body proportions and relative limb length are highly characteristic for most insect taxa. In case of the legs, limb length has mostly been discussed with regard to parameters of locomotor performance and, in particular cases, as an adaptation to environmental factors or to the mating system. Here, we compare three species of stick and leaf insects (Phasmatodea) that differ strongly in the length ratio between antennae and walking legs, with the antennae of Medauroidea extradentata being much shorter than its legs, nearly equal length of antennae and legs in Carausius morosus, and considerably longer antennae than front legs in Aretaon asperrimus. We show that that relative limb length is directly related to the near-range exploration effort, with complementary function of the antennae and front legs irrespective of their length ratio. Assuming that these inter-species differences hold for both sexes and all developmental stages, we further explore how relative limb length differs between sexes and how it changes throughout postembryonic development. We show that the pattern of limb-to-body proportions is species-characteristic despite sexual dimorphism, and find that the change in sexual dimorphism is strongest during the last two moults. Finally, we show that antennal growth rate is consistently higher than that of front legs, but differs categorically between the species investigated. Whereas antennal growth rate is constant in Carausius, the antennae grow exponentially in Medauroidea and with a sudden boost during the last moult in Aretaon.

Two new species of the genus Rhopalopsole (Plecoptera: Leuctridae) from China.

Two new species of the genus Rhopalopsole (Plecoptera: Leuctridae) are described from China: Rhopalopsole liui Chen, Wang Li, sp. nov. from from Municipality of Chongqing (former Sichuan Province) and Rhopalopsole yadonga Chen, Wang Li, sp. nov. from from the Tibet Autonomous Region. The new species are compared to similar congeners in the R. shaanxiensis and R. magnicerca groups, respectively.

Two new species of Oxyethira Eaton 1873 (Trichoptera: Hydroptilidae) from Roraima state, Brazil.

Two new species of Oxyethira from Brazil are described: Oxyethira (Dampfitrichia) jiquitaia sp. nov., and Oxyethira (Tanytrichia) librina sp. nov. Illustrations of male genitalia are provided with each description. These results increase the number of Oxyethira species in Brazil from 37 to 39.

Exploration of the homology among the muscles associated with the female genitalia of the three suborders of Psocodea (Insecta).

By using µCT technology, we reconstructed 3D models of the female genital structures and associated muscles of seven species from three suborders of Psocodea (free-living species only, formerly known as independent insect order "Psocoptera"). The homology of the female genital structures and associated muscles of different species is discussed. A total of 21 muscle groups were observed, and except for one muscle, all were homologized throughout the order. Moreover, some of the homologous muscles could be identified confidently in holometabolan insects. Using the muscles as landmarks, we discuss the homology of the ovipositor valves between Psocodea and other neopteran insects. Most importantly, the ovipositor of the suborder Trogiomorpha was identified to consist of the well-developed external valve (V3) plus a remnant of the dorsal valve (V2). We also examined the phylogenetic information included in the female genital muscles and found that certain muscles provide useful information and support deeper nodes (e.g., monophyly of the suborder Psocomorpha). The present study of female genital muscles not only helps us to better understand the phylogeny of Psocodea but also provides a solid foundation for research on muscle evolution.

Two new synonyms of Rhopalopsole basinigra Yang & Yang, 1995 (Plecoptera: Leuctridae) with taxonomic notes on Rhopalopsole tianmuana Sivec & Harper, 2008.

We present detailed evidence that Rhopalopsole trichotoma Yang & Du, 2021 and R. wulingensis Sivec & Shimizu, 2008, are junior synonyms of R. basinigra Yang & Yang, 1995 via a comparative morphological evaluation. We also present a comparison for R. basiningra from Guizhou Province, southwestern China, to understand the reliable characters of this species. New color images of male characters of R. tianmuana with some variations are provided.

Two new species of Amphinemura (Plecoptera: Nemouridae) from Taiwan Province on the southeast coast of China.

Two new species of Amphinemura were described and illustrated from Taiwan Province of China: Amphinemura reni Du Zhao, sp. nov. and Amphinemura triquetra Zhao Du, sp. nov. The morphological characteristics of the new species were compared to related taxa. Additionally, a key to the adult males of Amphinemura known from Taiwan Province of China was given in this paper.

A new species of Loneura Navás and taxonomic update of L. boliviana Williner and L. meridionalis García Aldrete (Psocodea: Psocomorpha: Ptiloneuridae).

A new species of Loneura named as Loneura willineri n. sp. is described from Bolivia, and and variation in the venation of the fore and hindwings discussed. Both sexes of Loneura meridionalis García Aldrete are described, inclulding variation in the fore and hindwing venation, and this species is withdrawn from synonymy with Loneura boliviana, a species now considered a nomen dubium.

A new species of Amphinemura (Plecoptera: Nemouridae) from Vietnam, with additions to A. bimaculata Li, Wang and Yang, 2016.

A new species of the Amphinemurinae genus Amphinemura, A. similibimaculata sp. nov. is described from central Vietnam. The new species is compared with related congeneric Amphinemura bimaculata Li, Wang and Yang, 2016. Taxonomic notes on A. bimaculata are also presented.

Penetration mechanics of elongated female and male genitalia of earwigs.

We unveiled the penile penetration mechanics of two earwig species, Echinosoma horridum, whose intromittent organ, termed virga, is extraordinarily long, and E. denticulatum, whose virga is conversely short. We characterised configuration, geometry, material and bending stiffness for both virga and spermatheca. The short virga of E. denticulatum has a material gradient with the stiffer base, whereas the long virga of E. horridum and the spermathecae of both species are homogeneously sclerotised. The long virga of E. horridum has a lower bending stiffness than the spermatheca. The virga of E. denticulatum is overall less flexible than the spermatheca. We compared our results to a previous study on the penetration mechanics of elongated beetle genitalia. Based on the comparison, we hypothesised that the lower stiffness of the male intromittent organ comparing to the corresponding female structure is a universal prerequisite for the penetration mechanics of the elongated intromittent organ in insects.

A new species of Neoperla/ (Plecoptera: Perlidae) from Motuo County of Tibet, China and redescription of Neoperla perspicillata/ Zwick, 1980.

A new species of the stonefly genus Neoperla, N. bimaculata sp. nov. is described from Motuo County of Tibet of southwest China. The new species compared with related congeners. Based on strongly pigmented specimens, we redescribe the male adult of Neoperla perspicillata Zwick, 1980.

A New Species of Moth Lacewing from the Mesoamerican Genus Adamsiana Penny 1996 (Neuroptera: Ithonidae).

The poorly studied Mesoamerican genus Adamsiana Penny, 1996 (Neuroptera: Ithonidae) was considered monotypic for more than 20 years, containing only Adamsiana curoei Penny. However, a second species was recently discovered in the southern region of Guatemala and is described here as Adamsiana alux Ardila-Camacho, Castillo-Argaez & Martinez, sp. nov. A key to the Adamsiana species and a list of the extant New World Ithonidae species are provided. This work emphasizes the necessity for more studies about not only Ithonidae but also all entomological fauna in the Neotropics.

Early postembryogenic development of the subgenual organ complex in the stick insect Sipyloidea sipylus.

Stick insects have elaborate mechanosensory organs in their subgenual organ complex in the proximal tibia, particularly the distal organ with scolopidial sensilla in linear arrangement. For early postembryonic developmental stages of Sipyloidea sipylus (Phasmatodea: Necrosciinae), the neuroanatomy of the scolopidial organs in the subgenual organ complex and the campaniform sensilla is documented by retrograde axonal tracing, and compared to the adult neuroanatomy. Already after hatching of the first larval instars are the sensory structures of subgenual organ and distal organ as well as tibial campaniform sensilla differentiated. In the distal organ, the full set of sensilla is shown in all larval stages examined. This finding indicates that the sensory organs differentiate during embryogenesis, and are already functional by the time of hatching. The constancy of distal organ sensilla over postembryonic stages allows investigation of the representative number of sensilla in adult animals as well as in larval instars. Some anatomical changes occur by postembryogenic length increase of the distal organ, and grouping of the anterior subgenual sensilla. The embryonic development of scolopidial sensilla is similar for auditory sensilla in hemimetabolous Orthoptera (locusts, bushcrickets, crickets) where tympanal membranes develop during postembryogenic stages, conferring a successive gain of sensitivity with larval moults.

A remarkable new genus and species of Nemourinae (Plecoptera, Nemouridae) from Sichuan, China, with systematic notes on the related genera.

A remarkable new genus and species of Nemourinae, Sinonemura balangshana gen. et sp. n., is described from Balang Mountains, Sichuan, southwestern China. The description is based on morphology and molecular data. The Nemourinae genera related to the new taxon are re-evaluated on the basis of comparative functional morphology of male epiproct. Notes on the Asian distribution of the Nemourinae are also given.

Viviparity in Two Closely Related Epizoic Dermapterans Relies on Disparate Modifications of Reproductive Systems and Embryogenesis.

Nutritional modes operating during embryonic/larval development of viviparous species range from "pure" lecitothrophy in which embryos rely solely on reserve materials (yolk spheres, lipid droplets, and glycogen particles) accumulated in the egg cytoplasm to matrotrophy in which embryos are continuously supplied with nutrients from a parental organism. Interestingly, a wide spectrum of diverse "mixed" modes employed in the embryo nourishment have also been described among viviparous species. Here, we summarize results of histochemical, ultrastructural, and biochemical analyses of reproductive systems as well as developing embryos of two closely related viviparous species of earwigs (Dermaptera), Hemimerus talpoides and Arixenia esau. These analyses clearly indicate that morphological as well as physiological modifications (adaptations) supporting viviparity and matrotrophy in Hemimerus and Arixenia, with the exception of a complex biphasic respiration, are markedly different. Most importantly, Hemimerus embryos complete their development inside terminal (largest) ovarian follicles, whereas Arixenia embryos, after initial developmental stages, are transferred to highly modified lateral oviducts, that is the uterus, where they develop until the release (birth) of larvae. The obtained results strongly suggest that viviparity in hemimerids and arixeniids had evolved independently and might therefore serve as an example of evolutionary parallelism as well as remarkable functional plasticity of insect reproduction and embryonic development.

Ecological gradients drive insect wing loss and speciation: The role of the alpine treeline.

Alpine ecosystems are frequently characterized by an abundance of wing-reduced insect species, but the drivers of this biodiversity remain poorly understood. Insect wing reduction in these environments has variously been attributed to altitude, temperature, isolation, habitat stability or decreased habitat size. We used fine-scale ecotypic and genomic analyses, along with broad-scale distributional analyses of ecotypes, to unravel the ecological drivers of wing reduction in the wing-dimorphic stonefly Zelandoperla fenestrata complex. Altitudinal transects within populations revealed dramatic wing reduction over very fine spatial scales, tightly linked to the alpine treeline. Broad biogeographical analyses confirm that the treeline has a much stronger effect on these ecotype distributions than altitude per se. Molecular analyses revealed parallel genomic divergence between vestigial-winged (high altitude) and full-winged (low altitude) ecotypes across distinct streams. These data thus highlight the role of the alpine treeline as a key driver of rapid speciation, providing a new model for ecological diversification along exposure gradients.

Morphogenesis of serial abdominal outgrowths during development of the viviparous dermapteran, Arixenia esau (Insecta, Dermaptera).

The embryos and first instar larvae of the epizoic earwig, Arixenia esau, develop sequentially in two different compartments of the female reproductive system, that is ovarian follicles and the lateral oviducts (the uterus). Here we show that the second (intrauterine) phase of development consists of three physiologically disparate stages: early embryos (before dorsal closure, surrounded by an egg envelope), late embryos (after dorsal closure, surrounded by an egg envelope) and the first instar larvae (after "hatching" from an egg envelope). Early and late embryos float in the fluid filling the uterus, whereas the first instar larvae develop attached to the uterus wall. Our analyses revealed also that in Arixenia serial multilobed outgrowths develop on dorso-lateral aspects of all abdominal segments. At the onset of the third developmental stage and after liberation from an egg envelope, these outgrowths (or more precisely their lobes) adhere to the epithelium lining the uterus, forming a series of small contact sites, where the mother and embryo tissues are separated only by a thin, presumably permeable, embryonic cuticle. We suggest that all these contact sites collectively constitute a dispersed placenta-like organ involved in the nourishment of the embryo.

Leptoceridae (Insecta, Trichoptera) from Serra da Canastra Mountain Range, Southeast Brazil: Diversity, Distribution, and Description of Two New Species.

The Brazilian savanna, also known as Cerrado, is one of the world's biodiversity hotspots, covering about 22% of the country. Nevertheless, this region has been suffering an accelerated process of degradation due to the agribusiness expansion. This study contributes to increasing knowledge on the Leptoceridae fauna of the Serra da Canastra Mountains at the Minas Gerais State, Brazil. Herein, we illustrate and describe two new species of Leptoceridae, Notalina franciscana sp. nov. and Oecetis catagua sp. nov., and provide additional data on the richness and distribution of Leptoceridae species in the Parque Nacional da Serra da Canastra and surrounding areas. Furthermore, ten species are firstly recorded from Minas Gerais State, increasing the number of Leptoceridae species recorded in the state from 30 to 42.