Wingless Insects Research Articles

Juvenile hormone signaling is indispensable for late embryogenesis in ametabolous and hemimetabolous insects

BackgroundJuvenile hormone (JH) is an insect-exclusive hormone involved in regulating diverse aspects of insect physiology, and the evolution of its diverse function is widely interesting. Studying embryogenesis in basal wingless insects is important to understand the functional evolution of JH; however, experimental studies in this regard are scarce. In this study, we conducted CRISPR/Cas9-mediated knockout (KO) of genes involved in JH biosynthesis and signaling cascades in the ametabolous firebrat, Thermobia domestica. Additionally, we investigated whether the primitive action of JH is conserved in the hemimetabolous cricket, Gryllus bimaculatus.ResultsWe observed that KO of JHAMT, CYP15A1, Met, and Kr-h1 resulted in embryonic lethality in T. domestica. Deprivation of JH or JH signaling arrested the progression of extraembryonic fluid resorption after dorsal closure and hatching, which is consistent with the gene expression pattern showing high Kr-h1 expression in the late embryos of T. domestica. The embryos deficient in JH signaling displayed wrinkled and weak legs. Comparative transcriptome analysis revealed that JH signaling promotes embryonic leg maturation through inducing energy supply and muscle activity, as validated by transmission electron microscopy (TEM). In addition, JH signaling exhibited similar embryonic effects in G. bimaculatus.ConclusionsThis study reveals the indispensable role of JH signaling in facilitating the maturation of terminal tissues during late embryogenesis, as demonstrated by the regulation of leg development, in ametabolous and hemimetabolous insects. These findings further indicate that the embryonic functions of JH evolved earlier than its anti-metamorphic functions during postembryonic development.

Evolutionary origin and distribution of leucine-rich repeat-containing G protein-coupled receptors in insects

Leucine-rich repeat-containing G protein-coupled receptors (LGRs) are crucial for animal growth and development. They were categorized into four types (A, B, C1, and C2) based on their sequence and domain structures. Despite the widespread distribution of LGRs across bilaterians, a thorough investigation of their distribution and evolutionary history remains elusive. Recent studies insect LGRs, especially the emergence of type C2 LGRs in various hemimetabolous insects, had prompted our study to address these problems. Initially, we traced the origins of LGRs by exploiting data from 99 species spanning 11 metazoan phyla, and discovered that type A and B LGRs originated from sponges, while type C LGRs originated from cnidarians. Subsequently, through comprehensive genomic and transcriptomic analyses across 565 species across 25 orders of insects, we found that both type A and C1 LGRs divided into two gene clusters. These clusters can be traced back to basal Insecta and an early ancestor of the Arthropoda, respectively. Furthermore, the absence of type B LGRs in wingless insects suggests a role in wing development, while the absence of type C2 LGRs in holometabolous insects hints at novel functions unrelated to insect metamorphosis. According to the origin of LGRs and the investigation of LGRs in insects, we speculated that type A and B LGRs appeared first among four types of LGRs, type A evolved into type C LGRs later, and type A and C1 LGRs independently duplicated during the evolutionary process. This study provides a more comprehensive view of the evolution of LGR genes than previously available, and sheds light on the evolutionary history and significance of LGRs in insect biology.

Wax "Tails" Enable Planthopper Nymphs to Self-Right Midair and Land on Their Feet.

The striking appearance of wax 'tails'-posterior wax projections on planthopper nymphs-has captivated entomologists and naturalists alike. Despite their intriguing presence, the functional roles of these formations remain largely unexplored. This study leverages high-speed imaging to uncover the biomechanical implications of wax structures in the aerial dynamics of planthopper nymphs (Ricania sp.). We quantitatively demonstrate that removing wax tails significantly increases body rotations during jumps. Specifically, nymphs without wax undergo continuous rotations, averaging 4.2 ± 1.8 per jump, in contrast to wax-intact nymphs, who do not complete a full rotation, averaging only 0.7 ± 0.2 per jump. This along with significant reductions in angular and translational velocity from takeoff to landing suggest that aerodynamic drag forces on wax structures effectively counteract rotation. These stark differences in body rotation correlate with landing success: Nymphs with wax intact achieve a near perfect landing rate of 98.5%, while those without wax manage only a 35.5% success rate. Jump trajectory analysis reveals that wax-intact jumps transition from parabolic to asymmetric shapes at higher takeoff velocities and show a significantly greater reduction in velocity from takeoff to landing compared to wax-removed jumps, demonstrating how wax structures help nymphs achieve more stable and controlled descents. Our findings confirm the aerodynamic self-righting functionality of wax tails in stabilizing planthopper nymph landings, advancing our understanding of the complex relationship between wax morphology and aerial maneuverability, with broader implications for wingless insect aerial adaptations and bioinspired robotics.

NERVOUS SYSTEM OF A PIGEON SLENDER LOUSE COLUMBICOLA COLUMBAE (PHTHIRAPTERA: ISCHNOCERA)

The nervous system of a pigeon slender louse, Columbicola columbae, has been studied in greater detail. C. columbae is a permanent, obligatory, and wingless insect. The central nervous system consists of a series of ganglia that supply nerves to successive body segments. The three main ganglia in the head, protocerebrum, deutocerebrum, and tritocerebrum, are fused to form the brain or supraesophageal ganglion. The nervous system of the pigeon slender louse, C. columbae (Phthiraptera: Ischnocera), consists of a U-shaped compact cerebral ganglion, subesophageal ganglion, and three thoracic ganglia. The ventral nerve cord and subesophageal ganglion are also located in the head. The three thoracic ganglions are placed in pro-, meso-, and meta-thorax. The frontal ganglion is situated anterior to the brain at a considerable distance.. KEYWORDS :Brain, Columbicola columbae, Nervous system, Pigeon slender louse

On some silverfish taxa from the Cape York region of northern Australia (Zygentoma: Lepismatidae: Ctenolepismatinae)

ilverfish (Zygentoma) are an ancient, but poorly studied group of primitively wingless insects. Species definition based on morphology alone is complicated by variations arising from the continuous moulting, even after reaching sexual maturity. In recent studies the use of molecular data has supported the identification of consistent morphological character traits, however only a small fraction of the species have been investigated. Eight species of the large silverfish genus Acrotelsella, collected by the first author from the Cape York Peninsula of northeastern Australia, are described and named as new species. Molecular data (COI and 28S) are presented that reinforces previous suggestions that the genus will eventually need to be split into two.

MAPHIS—Measuring arthropod phenotypes using hierarchical image segmentations

Abstract Animal phenotypic traits are utilised in a variety of studies. Often the traits are measured from images. The processing of a large number of images can be challenging; nevertheless, image analytical applications, based on neural networks, can be an effective tool in automatic trait collection. Our aim was to develop a stand‐alone application to effectively segment an arthropod from an image and to recognise individual body parts: namely, head, thorax (or prosoma), abdomen and four pairs of appendages. It is based on convolutional neural network with U‐Net architecture trained on more than a thousand images showing dorsal views of arthropods (mainly of wingless insects and spiders). The segmentation model gave very good results, with the automatically generated segmentation masks usually requiring only slight manual adjustments. The application, named MAPHIS, can further (1) organise and preprocess the images; (2) adjust segmentation masks using a simple graphical editor; and (3) calculate various size, shape, colouration and pattern measures for each body part organised in a hierarchical manner. In addition, a special plug‐in function can align body profiles of selected individuals to match a median profile and enable comparison among groups. The usability of the application is shown in three practical examples. The application can be used in a variety of fields where measures of phenotypic diversity are required, such as taxonomy, ecology and evolution (e.g. mimetic similarity). Currently, the application is limited to arthropods, but it can be easily extended to other animal taxa.

Resilin in Insect Flight Systems

AbstractCompared to wingless insects, pterygote insects profit from numerous wing‐related benefits including a wider distribution range, the exploitation of various food resources and the escape from water‐ or land‐confined predators. In order to maintain the wings´ functionality, the wing design and resistance to material fatigue are of key importance. This is even more essential for survival when considering that wings are used for millions of wing beat cycles but cannot be repaired and do not contain inner muscles so that their aerodynamic performance is mainly based on passive, structure‐based wing deformations. One of the components serving this purpose is the endowment of certain wing components with the elastomeric protein resilin building stable and complex material composites with the tanned cuticle. Resilin endows the respective structures with, e.g., higher flexibility and compliance and enables elastic energy storage. In this study, the occurrence of resilin in the insect flight system is reviewed based on previous studies of several insect orders including Odonata, Orthoptera, Hymenoptera, Coleoptera, Dermaptera, and Diptera, and the function of resilin is discussed with reference to the respective structures.

Development and Reproduction of a Japanese Strain of Ctenolepisma calvum (Ritter, 1910) at Room Temperature.

Ctenolepisma calvum (Ritter, 1910) (Zygentoma: Lepismatidae) is a primitive wingless insect that causes damage to paper, and it is regarded as a pest of collections in museums, archives, and libraries. This species was recently discovered in Japan for the first time and may have already spread over large areas of Japan, but, currently, no information is available on the biological characteristics of C. calvum in Japan. In this study, we observed the processes of development and reproduction of C. calvum found in Japan at room temperature. Oviposition was observed from April to November, with a peak in early June. The average egg period was 56.9 days at average temperatures above 24.0 °C, and was 72.4 days at average temperatures below 24.0 °C. The 1st, 2nd, and 3rd instars lasted 4.7 days, 13.2 days, and 26.1 days on average, respectively, at average temperatures above 22.0 °C. Average instar periods were 23-28 days in 4th-7th instars and tended to increase in later instars. Instar periods also increased when the average temperature was 22.0 °C or lower. In individual rearing, the longest-living individual lived for approximately two years, up to the 15th instar. The head width grew at an approximate ratio of 1.1 per molt. First oviposition occurred at the 10th or 11th instar. Individually observed females oviposited once or twice a year, laying 6-16 eggs at one time, but females at least two years old laid 78.2 eggs per year on average in a mass-culture cage. Through this study, only females were found, and the mature females produced their progenies parthenogenetically.

Revisiting the formation of midgut epithelium in Zygentoma (Insecta) from a developmental study of the firebrat Thermobia domestica (Packard, 1873) (Lepismatidae)

Zygentoma is an order of wingless insects, representing the sister group of Pterygota and constituting Dicondylia together with Pterygota. Contrasting views exist regarding midgut epithelium formation in Zygentoma. According to some reports, in Zygentoma, the midgut epithelium is entirely derived from yolk cells as in other wingless orders; however, according to other reports, the midgut epithelium is of dual origin in Zygentoma, similar to that in Palaeoptera of Pterygota, i.e., the anterior and posterior midgut epithelia are stomodaeal and proctodaeal respectively, whereas the middle part of the midgut originates from yolk cells. Aiming to provide a sound basis to evaluate the true image of midgut epithelium formation in Zygentoma, we examined the formation of the midgut epithelium in detail in Thermobia domestica, and concluded that the midgut epithelium is exclusively derived from yolk cells in Zygentoma, without the stomodaeal and proctodaeal elements involved in its formation. The participation of the anlagen differentiated at or around the stomodaeal and proctodaeal extremities in the formation of the midgut epithelium (bipolar formation) may be regarded as having first appeared not in Dicondylia but in Pterygota, of which the major part is represented by Neoptera with the midgut epithelium formed through bipolar formation.

GIS BASED FREQUENCY DISTRIBUTION OF WINGLESS INSECTS INFESTING DOMESTIC POPULATION OF DOGS IN LAHORE, PUNJAB, PAKISTAN

The intent of the present study was conducted (January, 2020 to December, 2020) to estimate the prevalence of wingless insects in dog population and to identify the potential risks factors in selected areas of Lahore, Punjab, Pakistan. Results of the study revealed an overall prevalence of wingless insects 50.16% (1204/2400) in dogs of study area. Among various insects, fleas were found predominant (28.5%; 684/2400; P<0.05) as compared to lice (21.66%; 520/2400). Among the reported species 4 were wingless insects (Ctenocepahlides (Ct.) felis, Ct, canis, Linognathus setosus and Trichodectes canis). Breed, age and sex of host were not found to be related with risk factors (P>0.05) affecting incidence of wingless insects. These inscets effect different body parts differently as highest prvelance was noted at neck (15%) and back (11.5%) followed by abdomen (10%) and ear (7.4%).Prevelence of the ectoparasites was found to be 6.2% at forelegs and 4% in congenital area, 4% around the shoulder and 6% around the hind legs, however, tail area was minimally affected (1.2%). Prevelance of Ct. felis was noted to be 35.96, Ct. canis 68.43%, Linognathus setosus 21.1% and Trichodectes canis 78.8%. Town wise incidences were found to be highest in Shalimar Town (62.91%) followed by Wagha Town (59.58%), Aziz Bhatti(59.16%), Data Ganj Bakhsh (55.41%), Nishtar Town (46.66%), Samanabad (44.3%), Iqbal Town (43.6%), Ravi Town (42%), Lahore Cantonment (41.6%) and Gulberg (40.4%). Rate of ectoparasites infestation was observed higher (P<0.05) in stray dogs (80.36%) followed by guard dogs (63.7%), hunting dogs (37.5%) and pet dogs (35.66%). Amongst the different seasons, rainy season showed higher lice prevalence (26.1%; 157/600), fleas (37.8%; 227/600), as compared to summer season and spring. Though, summer season was found to be unfavourable for the increase of almost all these insects. Comparative results of blood profile revealed significant difference (P<0.05) between non-infested and infested dogs, in the sort of lower value of infested/infected animals. To control the prevelance and ultimate effect of wingless insects on dog population applicable a well planned and control startegies as well as extension facilities should be arranged. Pet owners should be provided awareness about the significance of controlling the wingless insects to keep the pets and owner healthy.

The utility of probability plotting in palaeobiology

AbstractWe introduce probability plotting for general use in the analysis of biological and palaeontological morphometric data. When applied to morphometric data, this analytical tool can yield palaeoecological, ontogenetic and demographic information. We have adapted and expanded a statistical tool used in reliability engineering, mechanical or electrical lifetime analysis, geochemistry and ore mineral analysis to analyse fossil populations. The technique is explained stepwise in a ‘user's manual’ format. Its power is to characterize statistical distribution types, to determine whether these distributions are skewed or truncated, and to resolve and quantify the components of mixed distributions, which in palaeontological data potentially represent morphological groupings such as cohorts or sexual dimorphs. The ease of application and simplicity are surprising. A probability plot of growth series data (e.g. skull lengths, femoral lengths, snout‐to‐vent lengths or other morphometrics) typically shows complex non‐normal distribution of the data because of multimodal composition. We show that these component modes, which are generally, but not always normal (i.e. Gaussian), can be resolved, and that they correspond to yearly age groups (cohorts), sexual dimorphs, pre‐ and post‐metamorphic forms, other ontogenetic stages or possibly mixed taxa. When these component size distributions are associated with an age group or sex, several important palaeoecological calculations become possible. These include growth curves, age and sex distributions, sexual dimorphism indices, etc. We demonstrate the biological and palaeontological utility of probability plots with several examples including a Permian wingless insect, an extant unionid bivalve, a Triassic metoposaurid amphibian, an extant lizard, and a Triassic theropod dinosaur.

Possible role of Lipoptena fortisetosa (Diptera: Hippoboscidae) as a potential vector for Theileria spp. in captive Eld's deer in Khao Kheow open zoo, Thailand

Eld's deer (Rucervus eldii thamin) is an endangered species endemic to South Asia. Various ectoparasites (hematophagous insects and ticks) and blood parasites (e.g., piroplasms such as Babesia and Theileria) have been reported in this deer. Deer keds of the genus Lipoptena (L.) are wingless hematophagous insects acting as ectoparasites and potential vectors, thereby transmitting diseases to animals and humans. Many Lipoptena species have been reported, including L. fortisetosa; the latter may be a potential vector of several pathogens such as Babesia spp. and Theileria spp. However, the available data regarding Lipoptena in domestic animals and wildlife in Thailand is limited. The aim of this study was to investigate the presence of L. fortisetosa in Eld's deer as well as the role of this insect as a disease vector in Thailand by employing molecular analysis. A total of 91 wingless insects were collected and morphologically identified as L. fortisetosa. A partial fragment of the cytochrome c oxidase subunit I gene (COI) was amplified and successfully sequenced from twelve insects, and the COI nucleotide Basic Local Alignment Search Tool results revealed a 94.28%−94.45% identity to L. fortisetosa (accession number: OL850869/China). The undertaken phylogenetic analysis revealed that the L. fortisetosa samples from Thailand belong to a clade that is distinct from the previously deposited (in GenBank®) L. fortisetosa. As far as the pathogen detection is concerned, 46.2% (42/91) of the deer keds were positive for Theileria, while no Lipoptena was found to be positive for Babesia. Twenty-one sequences of Theileria were obtained and exhibited a 98.84%−100% identity to the Theileria sp. from several hosts. The phylogenetic analysis of Theileria revealed that Theileria capreoli and Theileria cervi were present in our L. fortisetosa samples. It can be implied that L. fortisetosa may serve as a vector of Theileria spp. in the Eld's deers of Thailand. We believe that the particular open zoo (from where the sampling took place) should implement preventive and control strategies for deer keds, other vectors, and vector-borne diseases.

Experimental infection of Aedes (Stegomyia) albopictus and Culex pipiens mosquitoes with Bluetongue virus.

Bluetongue disease (BT), caused by Bluetongue virus (BTV), infects wild and domestic ruminants, causing severe economic damage in the cattle and sheep industry. Proven vectors of BTV are biting midges belonging to the Culicoides genus, but other arthropods are considered potential vectors, such as ticks, mosquitoes, wingless flies, and sand flies. The present study represents the first attempt to evaluate the vectorial capacity of Culex pipiens and Aedes albopictus for BTV. Mosquitoes were artificially fed with blood containing BTV serotype 1. Infection, dissemination and transmission rates were evaluated at 0, 3, 7, 14 and 21 days after an infected blood meal. Viral RNA was only detected up to 3 days post infection in the bodies of both species. This study indicates that the two Italian populations of Cx. pipiens and Ae. albopictus are not susceptible to BTV infection.

Postembryonic development of the tracheal system of beetles in the context of aptery and adaptations towards an arid environment.

The tracheal system comprises one of the major adaptations of insects towards a terrestrial lifestyle. Many aspects such as the modifications towards wing reduction or a life in an arid climate are still poorly understood. To address these issues, we performed the first three-dimensional morphometric analyses of the tracheal system of a wingless insect, the desert beetle Gonopus tibialis and compared it with a flying beetle (Tenebrio molitor). Our results clearly show that the reduction of the flight apparatus has severe consequences for the tracheal system. This includes the reduction of the tracheal density, the relative volume of the trachea, the volume of the respective spiracles and the complete loss of individual tracheae. At the same time, the reduction of wings in the desert beetle allows modifications of the tracheal system that would be impossible in an animal with a functional flight apparatus such as the formation of a subelytral cavity as a part of the tracheal system, the strong elongation of the digestive tract including its tracheal system or the respiration through a single spiracle. Finally, we addressed when these modifications of the tracheal system take place during the development of the studied beetles. We can clearly show that they develop during pupation while the larvae of both species are almost identical in their tracheal system and body shape.

Abdominal serial homologues of wings in Paleozoic insects

The Late Paleozoic acquisition of wings in insects represents one of the key steps in arthropod evolution. While the origin of wings has been a contentious matter for nearly two centuries, recent evolutionary developmental studies suggest either the participation of both tergal and pleural tissues in the formation of wings1 or wings originated from exites of the most proximal leg podite incorporated into the insect body wall.2 The so-called "dual hypothesis" for wing origins finds support from studies of embryology, evo-devo, and genomics, although the degree of the presumed contribution from tergal and pleural tissues differ.3-6 Ohde etal.,7 confirmed a major role for tergal tissue in the formation of the cricket wing and suggested that "wings evolved from the pre-existing lateral terga of a wingless insect ancestor." Additional work has focused on identifying partial serially homologous structures of wings on the prothorax8,9 and abdominal segments.10 Thus, several studies have suggested that the prothoracic horns in scarab beetles,9 gin traps of tenebrionid and scarab beetle pupae,11,12 or abdominal tracheal gills of mayfly larvae1,13 evolved from serial homologues of wings. Here, we present critical information from abdominal lateral outgrowths (flaps) of Paleozoic palaeodictyopteran larvae, which show comparable structure to thoracic wings, consisting of cordate lateral outgrowths antero-basally hinged by muscle attachments. These flaps therefore most likely represent wing serial homologues. The presence of these paired outgrowths on abdominal segments I-IX in early diverging Pterygota likely corresponds to crustacean epipods14,15 and resembles a hypothesized ancestral body plan of a "protopterygote" model.

A hemimetabolous wing development suggests the wing origin from lateral tergum of a wingless ancestor

The origin and evolution of the novel insect wing remain enigmatic after a century-long discussion. The mechanism of wing development in hemimetabolous insects, in which the first functional wings evolved, is key to understand where and how insect wings evolutionarily originate. This study explored the developmental origin and the postembryonic dramatic growth of wings in the cricket Gryllus bimaculatus. We find that the lateral tergal margin, which is homologous between apterygote and pterygote insects, comprises a growth organizer to expand the body wall to form adult wing blades in Gryllus. We also find that Wnt, Fat-Dachsous, and Hippo pathways are involved in the disproportional growth of Gryllus wings. These data provide insights into where and how insect wings originate. Wings evolved from the pre-existing lateral terga of a wingless insect ancestor, and the reactivation or redeployment of Wnt/Fat-Dachsous/Hippo-mediated feed-forward circuit might have expanded the lateral terga.

Structural organization of xanthine crystals in the median ocellus of a member of the ancestral insect group Archaeognatha

Biogenic purine crystals function in vision as mirrors, multilayer reflectors and light scatterers. We investigated a light sensory organ in a primarily wingless insect, the jumping bristletail Lepismachilis rozsypali (Archaeognatha), an ancestral group. The visual system of this animal comprises two compound eyes, two lateral ocelli, and a median ocellus, which is located on the front of the head, pointing downwards to the ground surface.We determined that the median ocellus contains crystals of xanthine, and we obtained insights into their function. To date, xanthine biocrystals have only been found in the Archaeognatha. We performed a structural analysis, using reflection light microscopy, cryo-FIB-SEM, microCT and cryo-SEM. The xanthine crystals cover the bottom of a bowl-shaped volume in the median ocellus, in analogy to a tapetum, and reflect photons to light-sensitive receptors that are spread in the volume without apparent order or preferential orientation. We infer that the median ocellus operates as an irregular multifocal reflector, which is not capable of forming images. A possible function of this organ is to improve photon capture, and by so doing assess distances from the ground surface when jumping by determining changes in the intensity and contrast of the incident light.

Phoresy Involving Insects as Riders or Rides: Life History, Embarkation, and Disembarkation

AbstractThe ability to disperse is vital for all organisms, but especially for those whose habitats deteriorate, necessitating relocation to better feeding or breeding sites. Phoresy is assisted dispersal in which one organism uses another as its vehicle. In this review, phoresy will be largely restricted to cases wherein the rider is not parasitic on the vehicular stage used for dispersal, and in which insects are riders or rides. Phoresy occurs in organisms with limited mobility, and in secondarily brachypterous or completely wingless insects. Intraspecific phoresy also occurs. Generally, immature stages resistant to environmental assaults, such as dauers in nematodes, triungulins in beetles, or deutonymphs in mites, undertake phoresy. A size differential between rider and rider enables several conspecifics to board the same vehicle, likely ensuring mating opportunities at the destination. Riders may have special attachment devices or adhesive secretions to ensure safety during travel. Life cycles of riders and vehicles are often synchronized, and when phoresy is mutualistic, vehicles may also employ chemical tactics to achieve this synchrony. Chemical cues help to identify the appropriate vehicle, for disembarkation, or to lure the vehicle towards riders. Most riders prefer female vehicles, possibly to ensure access to rich nesting sites. Density-dependent selection may operate to limit the number of riders on a vehicle and phoresy may also facilitate sexual selection among riders. Whether vehicles can avoid phoresy is barely understood. This review attempts to place phoresy within a broad evolutionary context and points out the paucity of knowledge in certain research areas.

Old origin for an European‐African amphitropical disjunction pattern: New insights from a case study on wingless darkling beetles

AbstractAimThe origin of the amphitropic Mediterranean Basin and southern African disjunction (European–African amphitropical disjunction; EAAD) pattern is generally attributed to recent dispersal events. However, our knowledge is limited because the origin of the EAAD pattern has been almost exclusively studied in plants. Here, we investigate the origin of this wide‐ranging disjunction pattern in a group of wingless insects, consisting of two major clades, both of which have EAAD distributions.LocationSub‐Saharan Africa and Mediterranean region.TaxonTribe Dendarini (Coleoptera: Tenebrionidae).MethodsWe reconstructed a dated molecular phylogeny of major lineages within Dendarini using maximum likelihood and Bayesian inference. The employed dataset included sequences of six genes (two mitochondrial and four nuclear fragments) generated for 72 species. To investigate the sequence and timing leading to present‐day wide‐ranging disjunction patterns, we conducted parametric historical biogeography analyses.ResultsThe dated phylogenetic framework supports the monophyly of all major Dendarini lineages and highlights the origin of the tribe in sub‐Saharan Africa during the Middle Eocene. From there, representatives of the two major lineages colonized the Mediterranean region at the Oligocene‐Miocene boundary, with one lineage first reaching North Africa, whilst the other reached southern Europe.Main conclusionsThe origin of the EAAD in Dendarini beetles is ancient and better explained by the progressive fragmentation of the pan‐African rainforest that started in the Early Eocene than by other scenarios. This and the increased aridification associated with the global long‐term cooling trend that took place at that time had a strong influence on the diversification and distribution of xerophilic organisms such as dendarine beetles. This challenges the understanding of the origin of EAAD patterns, highlighting that they do not only result from recent dispersal events between the Pliocene and Pleistocene.

Spermiogenic chromatin condensation patterning in several hexapods may involve phase separation dynamics by spinodal decomposition or microemulsion inversion (nucleation)

We have examined published transmission electron microscopy (TEM).photomicrographs of chromatin condensation patterning in developing sperm nuclei from five species of entognathous hexapods within the Classes Protura, Collembola, Diplura and five species of ancestral wingless insects in the Orders Archaeognatha and Zygentoma as well as in fifteen species of the winged insects. Each species reproduces by internal fertilization. Spatially quantitative analysis indicates that spermiogenic chromatin condensation patterning in several of these species may be due to spinodal decomposition (SD) or to microemulsion inversion (chromatin-in-nucleoplasm → nucleoplasm-in-chromatin), also known as nucleation (Nc). These are two different dynamic mechanisms of liquid–liquid phase separation (LLPS). They might either occur independently or co-exist during the chromatin condensation associated with insect spermiogenesis. For example, the chromatin condensation pattern such as that observed in transverse sections of developing sperm nuclei from the wingless insect Anurida maritima (Collembola) is: granules → fibers → lamellae (SD) → nucleation (Nc) → condensed nuclei. Similar transitions are also observed in other more recently evolved species within the Class Insecta. From the limited but comprehensive sample of entognathus and ectognathus hexapods analyzed here, it appears that LLPS of sperm chromatin during spermiogenesis has occurred quite pervasively within the subphylum Hexapoda, including insects.