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The forewing base structure of the orthopteran suborder Ensifera, a group of insects well known for their sound-producing behavior, was examined by using light microscopy, confocal laser scanning microscopy, and µCT. We detected significant novel characters, such as functional changes in the tpm9 muscle from the flexor to the extensor of the forewing that are likely associated with the acquisition of sound-producing behavior using forewings. Phylogenetic analysis based on the characters selected from the forewing base structure showed that the character system contains a strong phylogenetic signal supporting the monophyly of Ensifera, Tettigonioidea and Grylloidea as well as the sister-group relationship between Grylloidea and Gryllotalpoidea, but no apomorphies to resolve the deep phylogeny among superfamilies could be determined from this character system.

To elucidate the effect of Pleistocene climatic fluctuations on the historical distribution and geographical genetic structure of temperate Japanese species, we performed phylogeographical and demographic analyses using mitochondrial gene sequences obtained from the stonefly species Kamimuria tibialis, sampled from four main islands of the Japanese Archipelago (i.e. Hokkaido, Honshu, Shikoku and Kyushu) and Tsushima Island. We detected three main clades with distinct geographical distributions, including the Tsushima, Kyushu and Hokkaido–Honshu–Shikoku phylogroups. These groups were estimated to have diverged from one another 0.54–2.02 Mya, suggesting they have undergone several glacial cycles in different refugia. Our results showed that during the glacial epochs and with a fall in sea-level, gene flow was limited among Tsushima and Kyushu, and among Hokkaido and Honshu, probably because the straits between these islands are deep. The population in Kyushu and Shikoku, the southernmost islands, exhibited high genetic diversity, with two distinct haplotype lineages occurring sympatrically. These results suggest that the population division into multiple refugia and the existence of stable southern refugia have contributed to the high genetic diversity of the species in this region.

The influence of the continental lithosphere and its root (or keel) on the continental drift of Earth is a key element in the history of plate tectonics. Previous geodynamic studies of mantle flow suggested that the cratonic root is moderately mechanically coupled with the underlying mantle, and stable continental drift on Earth's timescales occurs when the effective viscosity contrast between the continental lithosphere and the underlying mantle is approximately 103. Both geodynamics and seismological studies indicate that mechanically weak mobile belts (i.e., orogenic or suture zones) that surround cratons may play a role in the longevity of the cratonic lithosphere over geologically long timescales (i.e., over 1,000 million years) because they act as a buffer region against the high-viscosity cratons. Low-viscosity asthenosphere, characterized by slow seismic velocities, reduces the basal drag force acting on the cratonic root, which may also contribute to the longevity of the cratonic lithosphere. ▪ The role of the continental lithosphere and its root on the continental drift is reviewed from recent geodynamic and seismological studies. ▪ The cratonic root is moderately mechanically coupled with the underlying mantle and deformed by mantle flow over geological timescales. ▪ Orogenic belts or suture zones that surround cratons act as a buffer to protect cratons and are essential for their longevity. ▪ Low-viscosity asthenosphere may reduce the basal drag acting on the cratonic root and also contribute to its stability and longevity.

The thoracic musculature of the insect order Psocodea has been examined in only a few species of a single suborder to date. In the present study, we examined the thoracic musculature of species selected from all three suborders of Psocodea to elucidate the ground plan of the order and to examine the phylogenetic utility of the character system. The sister-group relationship between the suborders Troctomorpha and Psocomorpha received support from two novel nonhomoplasious synapomorphies, although the support from other morphological characters for this relationship is ambiguous. The sister-group relationship between the infraorders Epipsocetae and Psocetae also received support from one nonhomoplasious synapomorphy, although no other morphological characters supporting this relationship have been identified to date. The present examination revealed the potential of thoracic muscle characters for estimating deep phylogeny, possibly including interordinal relationships.

The explosive eruption occurred at Hunga Tonga-Hunga Ha'apai on January 15, 2022, generating atmospheric waves, meteotsunamis, tsunamis, and seismic waves recorded worldwide. Energetic signals of seismic surface waves indicate multiple explosive events, but the detailed eruption episodes are hard to unravel from dispersive long-period surface waves. In this study, we performed the back-projection (BP) analysis using the teleseismic P-waves to clarify the sequence of multiple explosive eruptions. The BP results successfully detected the locations and timing of multiple explosions even though P-waves from each eruption were overlapped and masked by other seismic signals and noises. The results suggest that the series of explosive eruptions can be divided into two parts of eruption sequences. The first sequence started at 04:02 UTC, escalated into major explosions at 04:15 UTC and 200 -300 s afterward, and lasted at least until 04:35 UTC. The second sequence began about 4 hours later (08:26 UTC) and continued for about 6 -7 minutes, including a massive eruption at 08:31 UTC. The estimated origin times of each eruption episode determined in this study are consistent with successive arrivals of long-period surface wave trains at frequencies higher than 10 mHz. The time intervals of explosive eruptions in the first sequence suggested that the significant explosive eruptions intermittently occurred at around 270 -280 s intervals. This interval is nearly equivalent to the eigenfrequencies of the atmospheric resonant oscillation with the Earth at 3.7 mHz. Since the observed amplitude spectra of this eruption exhibit a conspicuous spectral peak at 3.7 mHz, our BP results indicate that the eruption cycle may be synchronized with the eigen-period of the atmosphere-lithosphere coupled mode. (c) 2022 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

info:eu-repo/semantics/publishedVersion

SUMMARYRadially anisotropic S-wave structures under the permanent seismic stations in Australia are reconstructed using multimode surface waves (SWs) and receiver functions (RFs) in a framework of the Bayesian inference. We have developed a fully nonlinear method of joint inversions incorporating P-RFs and multimode Rayleigh and Love waves, based on the transdimensional Hierarchical Bayesian formulation. The method allows us to estimate a probabilistic Earth model taking account of the complexity and uncertainty of Earth structure, by treating the model parameters and data errors as unknowns. The Parallel Tempering algorithm is employed for the effective parameter search based on the reversible-jump Markov Chain Monte Carlo method. The use of higher modes enables us to enhance the sensitivity to the depth below the continental asthenosphere. Synthetic experiments indicate the importance of higher mode SWs for the better recovery of radial anisotropy in the whole depth range of the upper mantle. The method is applied to five Global Seismographic Network stations in Australia. While the S-wave models in eastern Australia show shallow lithosphere–asthenosphere boundary (LAB) above 100 km depth, those in central and Western Australia exhibit both mid-lithosphere discontinuities (MLDs) and LAB. Also, seismic velocity jumps equivalent to the Lehmann discontinuity (L-D) are found in all seismic stations. The L-D under the Australian continents is found at around 200–300 km depth, depending on locations. Radial anisotropy in the depth range between LAB and L-D tends to show faster SH anomalies, which may indicate the effects of horizontal shear underneath the fast-moving Australian plate.

The Japanese species of the genus Strigiphilus Mjöberg, 1910 (Insecta: Phthiraptera: Philopteridae) are revised. Six species are recorded, including a new species belonging to the cursitans species-group: Strigiphilus stenocephalus new species, described from the type host Otus bakkamoena semitorques and based on specimens originally identified and reported by Uchida (1949) as Strigiphilus rostratus (Burmeister, 1838). A lectotype for Strigiphilus laticephalus (Uchida, 1949) (type host: Strix aluco yamadae) is designated and redescribed, and this louse species is synonymized under Strigiphilus cursor (Burmeister, 1838). Strigiphilus ceblebrachys (Denny, 1842), S. heterogenitalis Emerson & Elbel, 1957 and S. tuleskovi Balát, 1958 are recorded for the first time in Japan. Also, Strix uralensis and Otus sunia japonicus are recorded as new hosts for Strigiphilus heterogenitalis and S. tuleskovi respectively.

Cormopsocidae n. fam. of the psocid suborder Trogiomorpha was proposed for a fossil species from mid‐Cretaceous Burmese amber, Cormopsocus groehni n. gen. & n. sp. This family was estimated to be the sister group of all other trogiomorphan taxa, but the possibility of much deeper divergence (i.e. placement as a sister group of all Psocodea) could not be excluded. Cormopsocus groehni retains many plesiomorphic features, which will contribute greatly to elucidating the ancestral state of Psocodea.

By using mu CT 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. (C) 2022 Elsevier Ltd. All rights reserved.

The divergence date and ancestral distributional area of the psocid subfamily Speleketorinae, which includes taxa with reversed genitalia (female penis and male vagina of Afrotrogla and Neotrogla, tribe Sensitibillini), were estimated. The most basal divergence of the subfamily (between the North American Speleketor and the tribe Sensitibillini) was estimated to have occurred according to the separation between the North American continent and Gondwana, ca. 175 Ma. The most basal divergence of Sensitibillini (between African Afrotrogla + Sensitibilla and Brazilian Neotrogla) was estimated to have occurred according to the split of West Gondwana (separation between the African and South American continents), ca. 127 Ma. The biome of the ancestral distributional area of Sensitibillini (inland of West Gondwana) is believed to be arid to semi-arid, which might strengthen the reversed sexual selection and then facilitate the origin of preadaptive features related to the evolution of a female penis. All extant Sensitibillini species inhabit carbonatic caves, but geological evidence suggested independent shifts of these genera to the carbonatic cave habitat in the Tertiary/Quaternary.

Using broadband seismic waveforms recorded on the seafloor by more than 200 broadband ocean bottom seismometers, as well as those on land, we measured the phase speed dispersions of Love and Rayleigh waves up to the 4th higher mode to determine the three-dimensional radially anisotropic shear-wave speed structure in the upper mantle beneath the Pacific Ocean. The fastest anomalies at depths shallower than 100 km were located beneath southeast of the Shatsky Rise and strong radial anisotropy was located in the central Pacific at depths of 100–200 km. The isotropic shear-wave speed structures showed age dependence. From the age-bin-averaged shear-wave speed profiles and a half-space cooling model, we estimated the thermo-speed relationship for the Pacific Plate to construct a reference age-dependent shear-wave speed model, which was further used as an initial model for tomography iteration. Deviation maps in the Pacific Ocean from the reference model indicated that large negative residuals, which may be due to partial melting, anelasticity, and/or added heat from mantle plumes, were located along the ridges and beneath hotspots, and that large positive residuals were found beneath the northwestern Pacific Ocean. The use of an age-dependent reference model, as well as the incorporation of OBS data, greatly improves the accuracy of local phase speed estimates in tomography, as evident from a direct comparison with in situ array measurements.

The systematic placement of an enigmatic psocid family restricted to Africa, Lesneiidae, was estimated by using a multiple gene data set. The candidates for its close relatives are now classified under two different infraorders, the family Archipsocidae of the infraorder Archipsocetae or the families Elipsocidae/Mesopsocidae of the infraorder Homilopsocidea. The maximum likelihood and Bayesian analyses of the molecular data set strongly suggested that the Lesneiidae belongs to Homilopsocidea and forms a clade with Elipsocidae/Mesopsocidae/Eolachesillinae (Lachesillidae). However, the relationships among these (sub)families and Lesneiidae, including the monophyly of Elipsocidae and Mesopsocidae, were ambiguous or questionable, showing the necessity of further investigations for elucidating their relationships and validating the status of these families. Two species, L. johnsoni Yoshizawa & Lienhard, n. sp. and L. testudinata Yoshizawa & Lienhard, n. sp. (Psocodea: Lesneiidae), were described from South Africa. There appears to be a tight association between the reproductive biology and morphological specialization of this group.

Many male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in a higher paternity Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and female polyandry, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to analyze such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Our genetically explicit, individual-based computer simulations demonstrate that a positive correlation between donated gift size and the resultant paternity gain is a requisite for the co-occurrence of large-sized gifts and females’ competitive multiple mating for the gifts. When donation of gifts imposes monandry, exaggeration in nuptial gift size also occurs under the assumption that the last male monopolizes paternity. We also analyzed the causes and consequences of the evolution of a female persistence trait in trading of nuptial gifts, that is, double receptacles for nuptial gifts known to occur in an insect group with a “female penis” (Neotrogla spp.).

The Ontong Java Plateau (OJP), one of the largest oceanic plateaus located in the western Pacific Ocean, was first formed by a massive volcanism at 122 Ma, which had a major effect on the Earth's environments, including global climate change, oceanic anoxic events, and mass extinction of marine life. However, the cause of the volcanism remainscontroversial since the underground structure beneath the OJP has been poorly understood due to limited geophysical and geochemical data. To improve such situation, we conducted about 1.6-year long-term seafloor observation on the OJP and its vicinity. Using seismograms obtained by this observation as well as those from existing seismic stations, we obtained three dimensional radially anisotropic shear wave velocity structure beneath the OJP at depths down to 300 km.Obtained structure shows the following new features:(1) Beneath the Caroline Islands, in the north of the OJP, 1 % slow anomalies exist, which may be associated with the Caroline hotspot activity;(2) In the center of the OJP at depths between 70–130 km, about 2% fast anomalies, whose shear wave speed is about 4.45-4.55 km/s, exists.(3) The seismic structure clearly shows that the lithosphere–asthenosphere boundary (LAB) beneath the center of the OJP is located about 40 km deeper than that beneath the surrounding normal oceanic seafloor.Judging from our results and petrological/rheological constraints given by previous studies, we interpret that the LAB is deepened by dehydrated residual material from hot mantle plume underplating a pre-existing lithosphere during a formation of OJP.

Upper mantle structures under cratons have recently been investigated by many researchers using receiver functions and surface waves to clarify the nature of the Lithosphere-Asthenosphere Boundary (LAB) and Mid-Lithosphere Discontinuity (MLD). Majority of seismological studies of joint inversions using receiver functions and surface waves have employed dispersion curves of fundamental-mode only, but higher-mode information is essential for resolving the whole depth range of thick continental lithosphere (over 200 km) and its underlying asthenosphere.In this study, we reconstructed radially anisotropic S wave models including multiple discontinuities in the upper mantle under seismic stations in Australia, using multi-mode surface waves and receiver functions in the framework of the Bayesian inference. We employed a fully nonlinear method of joint inversions incorporating P-to-S receiver functions and multi-mode Rayleigh and Love waves, based on the trans-dimensional hierarchical Bayesian formulation. The method allows us to estimate a probabilistic Earth model taking account of the complexity and uncertainty of Earth structure, by treating the model parameters and data errors as unknowns. The Parallel Tempering algorithm is incorporated for the effective parameter search based on the reversible-jump Markov Chain Monte Carlo method.Multi-mode phase speed maps of surface waves developed by Yoshizawa (2014) are used to extract localized multi-mode dispersion curves. The use of higher-mode surface waves enables us to enhance the sensitivity to the depth below the continental asthenosphere, while the receiver functions allows us to better constrain the depths of discontinuities and velocity jumps. Synthetic experiments indicate the importance of higher-mode information for the better recovery of radial anisotropy in the whole depth range of the upper mantle.The method has been applied to Global Seismographic Network stations in Australia. While the S-wave models in eastern Australia show shallow LAB above 100 km depth, those in central and western Australia exhibit both MLD and LAB. Also, seismic velocity jumps equivalent to the Lehmann Discontinuity (LD) are found in all seismic stations in Australia. The LDs under the Australian continents are found at the depth of around 200 - 300 km, depending on locations. Radial anisotropy in the depth range between LAB and LD tends to show faster SH anomalies, which may indicate the effects of horizontal shear underneath the fast-moving Australian plate.

The insect order Psocodea is a diverse lineage comprising both parasitic (Phthiraptera) and nonparasitic members (Psocoptera). The extreme age and ecological diversity of the group may be associated with major genomic changes, such as base compositional biases expected to affect phylogenetic inference. Divergent morphology between parasitic and nonparasitic members has also obscured the origins of parasitism within the order. We conducted a phylogenomic analysis on the order Psocodea utilizing both transcriptome and genome sequencing to obtain a data set of 2370 orthologous genes. All phylogenomic analyses, including both concatenated and coalescent methods suggest a single origin of parasitism within the order Psocodea, resolving conflicting results from previous studies. This phylogeny allows us to propose a stable ordinal level classification scheme that retains significant taxonomic names present in historical scientific literature and reflects the evolution of the group as a whole. A dating analysis, with internal nodes calibrated by fossil evidence, suggests an origin of parasitism that predates the K-Pg boundary. Nucleotide compositional biases are detected in third and first codon positions and result in the anomalous placement of the Amphientometae as sister to Psocomorpha when all nucleotide sites are analyzed. Likelihood-mapping and quartet sampling methods demonstrate that base compositional biases can also have an effect on quartet-based methods.[Illumina; Phthiraptera; Psocoptera; quartet sampling; recoding methods.]

The gain of foldable wings is regarded as one of the key innovations enabling the present-day diversity of neopteran insects. Wing folding allows compact housing of the wings and shields the insect body from damage. Wing-fixing systems have evolved in some insects, probably to increase the durability of the shielding function by the wings. Bark lice (Psocodea) are known to possess a unique wing-to-wing repose coupling system, but a detailed morphological and evolutionary study of this system is lacking. In this study, we examined this repose coupling structure by SEM in 32 species including representatives of all three suborders of bark lice (Trogiomorpha, Troctomorpha and Psocomorpha). We concluded that the repose wing-coupling apparatus independently evolved twice within Psocodea. In Trogiomorpha, the apparatus is located on the subcostal vein of the forewing and is composed of elongated rib-like structures. In Troctomorpha and Psocomorpha, in contrast, the repose coupling structure is located on the radius vein of the forewing and is formed by a swollen vein. These morphological and developmental differences in the repose coupling structures also provide phylogenetic information at different systematic levels.

Many four-winged insects have mechanisms that unite the forewings and hindwings in a single plane. Such an in-flight wing coupling apparatus may improve flight performance in four-winged insects, but its structure is variable among different insect groups. The wings of bark lice (Insecta: Psocodea: "Psocoptera") also have an in-flight wing coupling apparatus, but to date, its morphology has not been studied in detail. In this study, we examined the wing-coupling structure in representative species of the three suborders of bark lice (Trogiomorpha, Troctomorpha, and Psocomorpha) and inferred its origin and transformation. We conclude that the main component of the psocodean wing coupling apparatus evolved once in the common ancestor via modification of cuticular structures at the apex of the forewing CuP vein. Morphological differences in components of the coupling structures are phylogenetically informative at the intraorder level and include an autapomorphy that characterizes Troctomorpha and a synapomorphy that supports a sister relationship between Troctomorpha and Psocomorpha.

The mitochondrial genome arrangement in the insect order Psocodea (booklice, barklice, and parasitic lice) is extremely variable. Genome organization ranges from the rearrangement of a few tRNAs and protein coding genes, through extensive tRNA and protein coding gene rearrangements, to subdivision into multiple mini-chromosomes. Evolution of the extremely modified mitochondrial genome in parasitic lice (Phthiraptera) has been the subject of several studies, but limited information is available regarding the mitochondrial genome organization of the more plesiomorphic, free-living Psocodea (formerly known as the "Psocoptera"). In particular, the ancestral state of the psocodean mitochondrial genome arrangement and the evolutionary pathway to the rearranged conditions are still unknown. In this study, we addressed mitochondrial evolutionary questions within the Psocodea by using mitochondrial genome sequences obtained from a wide range of Psocoptera, covering all three suborders. We identified seven types of mitochondrial genome arrangements in Psocoptera, including the first example in Psocodea of retention of the ancestral pancrustacean condition in Prionoglaris (Prionoglarididae). Two methods (condition-based parsimony reconstruction and common-interval genome distances) were applied to estimate the ancestral mitochondrial arrangement in Psocodea, and both provided concordant results. Specifically, the common ancestor of Psocodea retained the ancestral pancrustacean condition, and most of the gene arrangement types have originated independently from this ancestral condition. We also utilized the genomic data for phylogenetic estimation. The tree estimated from the mitochondrial genomic data was well resolved, strongly supported, and in agreement with previously estimated phylogenies. It also provided the first robust support for the family Prionoglarididae, as its monophyly was uncertain in previous morphological and molecular studies.

International audience; Palaeosiamoglaris hkamtiensis sp. nov. of the family Prionoglarididae is described from Hkamti amber (ca. 110 Ma), together with Psyllipsocus myanmarensis sp. nov., the third fossil psyllipsocid species described from from Noije Bum Hill (ca. 98 Ma). Based on the current observations of Palaeosiamoglaris hkamtiensis sp. nov., we discuss the synapomorphic characters proposed to define the tribe Siamoglaridini. Palaeosiamoglaris hkamtiensis sp. nov. is, inter alia, characterized by the maxillary palpomere 3 ca. 0.66 time as long as maxillary palpomere 4, the second anal vein concavity turned toward posterior wing margin in forewing, Sc reaching R1 nearly at middle of radial cell. The new species Psyllipsocus myanmarensis sp. nov. is, inter alia, characterized by antennae with 11 segments, the radial cell closed and six-angled, and the quadrangular pterostigma.

Understanding the complex heterogeneity of the continental lithosphere involves a wide variety of spatial scales and the synthesis of multiple classes of information. Seismic surface waves and multiply reflected body waves provide the main constraints on broad-scale structure, and bounds on the extent of the lithosphere-asthenosphere transition (LAT) can be found from the vertical gradients of S wavespeed. Information on finer-scale structures comes through body wave studies, including detailed seismic tomography and P-wave reflectivity extracted from stacked autocorrelograms of continuous component records. With the inclusion of deterministic large-scale structure and realistic medium-scale stochastic features fine-scale variations are subdued. The resulting multi-scale heterogeneity model for the Australian region gives a good representation of the character of observed seismograms and their geographic variations and matches the observations of P-wave reflectivity. P reflections in the 0.5–3.0 Hz band in the uppermost mantle suggest variations on vertical scales of a few hundred metres with amplitudes of the order of 1%. Interference of waves reflected or converted at sequences of such modest variations in physical properties produce relatively simple behaviour for lower frequencies, which can suggest simpler structures than are actually present. Vertical changes in the character of fine-scale heterogeneity can produce apparent discontinuities. In Central Australia a ‘mid-lithospheric discontinuity’ can be tracked via changes in frequency content of station reflectivity, with links to the broad-scale pattern of wavespeed gradients and, in particular, the gradients of radial anisotropy. Comparisons with xenolith results from southeastern Australia indicate a strong tie between geochemical stratification and P-wave reflectivity.

The Ontong Java Plateau in the western Pacific Ocean is the world’s largest oceanic plateau. It was formed 122 million years ago by a massive volcanic event that significantly affected Earth’s environment. The cause of the magmatic event remains controversial because the upper mantle structure beneath the plateau is poorly known. Here we use passive seismic data obtained through seafloor observations, alongside existing seismic data, to determine the three-dimensional radially anisotropic shear wave velocity to depths of up to 300 km. We find that the lithosphere–asthenosphere boundary is approximately 40 km deeper beneath the centre of the Ontong Java Plateau than beneath the surrounding seafloor. Based on our results and petrological and rheological constraints, we propose that the lithosphere–asthenosphere boundary has deepened as a result of underplating of dehydrated residual material beneath the pre-existing lithosphere during formation of the Ontong Java Plateau by a thermochemical mantle plume. The lithosphere beneath the Ontong Java Plateau is thickened by up to 40 km relative to the surrounding ocean floor which suggests it formed through the ascent of a mantle plume, according to analyses of seafloor seismic observations.

Morphological and behavioral characters are frequently examined for comparative studies. Unlike morphology, a single behavioral trait is difficult to subdivide as multiple characters, even when achieved by many evolutionary changes. Therefore, when similar behavioral traits evolved independently among closely related taxa, their distinction is difficult. Almost all members of the suborder Auchenorrhyncha (Insecta: Hemiptera) possess a jumping ability that uses metathoracic muscles, and this behavioral trait has been regarded as a synapomorphy. In this study, the anatomical observations of metathoracic muscles revealed that highly elaborated jumping ability was gained independently within the suborder, although the evolution of jumping ability might have been initiated at their common ancestor. Our results provide an example of identifying a true evolutionary pathway by dissecting a behavioral character into mechanical elements.

A new barklouse species belonging to the Trichadenotecnum s.str. clade (Psocodea: “Psocoptera”: Psocidae) was described from southeastern Brazil (Minas Gerais state). This locality is highly isolated (about 3400 km) from the closest known distributional range of this clade. It also represents the southernmost distributional record of Trichadenotecnum s.str. Phylogenetic placement of the species and biogeography of Neotropical Trichadenotecnum were briefly discussed. Trichadenotecnum punctipenne New, 1972 described from Brazil was synonymized with Trichadenotecnum pardus Badonnel, 1955.

Three species of chewing lice—Eucolpocephalum femorale (Piaget, 1880) (Amblycera: Menoponidae), Ibidoecus plataleae (Denny, 1842) (Ischnocera: Philopteridae) and Ardeicola plataleae (Linnaeus, 1758) (Ischnocera: Philopteridae)—are reported from the black-faced spoonbill (Platalea minor Temminck & Schlegel, 1849) in Japan. These three louse species are new records for this host. The black-faced spoonbill is classified as a “Critically Endangered” species in the IUCN Red List; although its species of lice are known to infest other species of spoonbills, these lice are endangered at the population level.

Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dataset comprising sequences of 2,395 single-copy, protein-coding genes for 193 samples of hemipteroid insects and outgroups. These analyses yield a well-supported phylogeny for hemipteroid insects. Monophyly of each of the three hemipteroid orders (Psocodea, Thysanoptera, and Hemiptera) is strongly supported, as are most relationships among suborders and families. Thysanoptera (thrips) is strongly supported as sister to Hemiptera. However, as in a recent large-scale analysis sampling all insect orders, trees from our data matrices support Psocodea (bark lice and parasitic lice) as the sister group to the holometabolous insects (those with complete metamorphosis). In contrast, four-cluster likelihood mapping of these data does not support this result. A molecular dating analysis using 23 fossil calibration points suggests hemipteroid insects began diversifying before the Carboniferous, over 365 million years ago. We also explore implications for understanding the timing of diversification, the evolution of morphological traits, and the evolution of mitochondrial genome organization. These results provide a phylogenetic framework for future studies of the group.

Although the great genital diversity of the barklouse genus Trichadenotecnum has been described in previous studies, the specific function of the genital structures during the copulation process has received less investigative attention. We reconstructed 3D-models of each structure of the male and female genitalia of Trichadenotecnum incognitum in copula and those of uncopulated male and female of Trichadenotecnum pseudomedium. By comparing the changes in male and female genital structures and related muscles in copulated and uncopulated states, the function of each genital structure can be described. During the copulation, we found that the female subgenital plate was hooked into the male body by the distal process on the male paraproct and was fixed by the male epiproct, hypandrium and phallosome. In addition, sexual coevolution was suggested by tightly contacting structures, that is, thorny male hypandrium and thickened membrane around the female spermapore plate. These results not only give us a new understanding copulation process of Trichadenotecnum, but also explain the reasons why genital structures are so divers in the genus.

In contrast to active continental rifting, which is caused by the thermal erosion of active upwelling plumes, passive continental rifting is attributed to the extensional stress acting on the continental lithosphere due to either the basal drag force exerted by the passive mantle flow under the lithosphere or the boundary forces at the continental margins. Although the extension of continental lithosphere by passive continental rifting may cause the breakup of continental margins, such as the separation of Zealandia from Gondwanaland at ca. 100 Ma and the Japanese islands from Eurasia at ca. 25 Ma, the dynamics have not been fully understood from geophysical and geological perspectives. Here, a series of numerical experiments of visco-elasto-plastic thermo-chemical convection with a free surface is performed using 2-D Cartesian models to investigate the localization of strain in the extending continental lithosphere, with an initial depth of 200 km, and the behavior of continental rifting and breakup. Our results demonstrate that when extension rates at the continental margin were set between 1 and 4 cm yr−1, the time taken for seafloor subsidence and subsequent continental breakup under extensional stress is less than a few million years. During continental rifting, high-shear zones (i.e., high-strain-rate zones) develop under the base of the deforming continental lithosphere, with a strain rate of the order of 10−13 s−1, which is approximately two orders larger than the typical value of the mantle interior. Due to accumulating strain along the base of the deforming lithosphere, excess strain is not concentrated on the shallowest part of the underlying mantle; hence, the spreading ridge that breaks the thick continental lithosphere emerges as a locally weak plate boundary on the Earth's surface. This characteristic is more remarkable as the activation volume of the sublithospheric mantle rock and the resultant viscosity are low under hydrous conditions.

Trichadenotecnum is one of the most diverse genera among the non-parasitic members of Psocodea (Insecta: "Psocoptera"). The genus shows a world-wide distribution (excluding the Australian Region, where only one introduced species is known) with its center of diversity in southern to eastern Asia. Several species groups had been proposed for this large genus based on morphology, but their validity and phylogenetic relationships are still unclear because of great morphological diversity in the genitalia, systematically the most relevant character. In this study, we estimated the molecular phylogeny of the Old World species of Trichadenotecnum based on extensive taxon sampling. As a result, the monophyly of morphology-based species groups was very strongly supported in most cases. However, two groups were recovered as non-monophyletic, which had been inadequately defined on the basis of plesiomorphies or convergences of genital characters. First, the monophyly of the sexpunctatum group was not supported because the medium group was found to be embedded within this group. The simpler genitalia observed in the medium group were considered to be derived from the more complicated genitalia present in the sexpunctatum group. Second, the monophyly of the majus group was not supported for two reasons: (1) It was divided into two distant clades which initially had been united on the basis of convergent similarities of the male genitalia. (2) Two species groups were revealed to be embedded within the main clade of the majus group; the initial separation of these groups had been based on reversals to the ancestral genital condition.

Two end-member scenarios have been proposed for the tectonic situation along the eastern margins of Gondwanaland before Zealandia was formed ca. 100 million years ago (Ma), namely: (1) A subduction zone located far from the eastern margin of Zealandia, wherein Zealandia may have separated from Gondwanaland by plume push of an active hotspot plume.; (2) A subduction zone located along the eastern margin of Gondwanaland, wherein Zealandia possibly separated from Gondwanaland via trench/subduction retreat. Assuming that the thermal structure of the deep mantle and source of hotspot plumes remained relatively stationary over the last hundred million years, major hotspot plumes with a large buoyancy flux did not exist under Zealandia; the eastern margins of Gondwanaland were far from two large low-shear-velocity provinces under the Africa–Atlantic and South Pacific regions. Herein, through numerical studies of three-dimensional global mantle convection, we examined the mantle convection and surface tectonic patterns at ~100 Ma. The present model considered the real configuration of Gondwanaland at the model surface to observe long-term variations of mantle convection and the resulting surface tectonic conditions. The results demonstrate that the extensive subduction zones developed preferentially along the eastern margin of Gondwanaland when the temperature anomaly of the lower mantle was primarily dominated by high-temperature regions under present-day Africa–Atlantic and South Pacific regions. The results of this study support one of the proposed hypotheses, where the breakup at the eastern margins of Gondwanaland at ~100 Ma occurred via trench/subduction retreat.

The cave-dwelling psocid tribe Sensitibillini ( Afrotrogla , Neotrogla and Sensitibilla ) is of special morphological and evolutionary interest because of its possession of reversed copulatory organs: i.e. females of Afrotrogla and Neotrogla have a penis-like organ. The female penis structure is highly variable among taxa, as is the case of the male penis in animals with normal copulatory organs. Here, we present the first molecular phylogeny of Sensitibillini and analyse the evolutionary pattern of their genitalia. Afrotrogla and Neotrogla did not form a monophyletic clade, and their female penis structures are significantly different, suggesting two independent origins of the female penis within Sensitibillini. In Neotrogla , the species that has a simple female penis is embedded among species that have an elaborate penis, and detailed structures of the female penis elaborations are in exact agreement among species, suggesting a secondary simplification of the female penis. A correlated evolutionary pattern between male and female genitalia was also detected. This coevolution of genitalia may suggest that sexual conflict or cryptic ‘male’ choice drove the diversity of the female penis, as is the case of male penile diversity in animals with conventional genitalia.

We report a functional switching valve within the female genitalia of the Brazilian cave insect Neotrogla. The valve complex is composed of two plate-like sclerites, a closure element, and in-and-outflow canals. Females have a penis-like intromittent organ to coercively anchor males and obtain voluminous semen. The semen is packed in a capsule, whose formation is initiated by seminal injection. It is not only used for fertilization but also consumed by the female as nutrition. The valve complex has two slots for insemination so that Neotrogla can continue mating while the first slot is occupied. In conjunction with the female penis, this switching valve is a morphological novelty enabling females to compete for seminal gifts in their nutrient-poor cave habitats through long copulation times and multiple seminal injections. The evolution of this switching valve may have been a prerequisite for the reversal of the intromittent organ in Neotrogla., eLife digest In dry caves of southeastern Brazil, live a group of insects named Neotrogla that are perhaps best known because the egg-producing females have penises while the sperm-producing males have vaginas. The sex roles of these Brazilian cave insects are also reversed: females compete over the males, who in turn are selective of their female partners. This sex role reversal likely evolved within Neotrogla because the males’ semen represents a rich and reliable source of energy within a nutrient-poor cave environment. When females are not using semen to fertilize their eggs, they consume it. Yet, while other animals show sex role reversal, Neotrogla species alone have reversed sexual organs. Neotrogla penises are spiky and may have evolved so that females can anchor themselves to males and then mate for prolonged periods. This would allow the females to stock up on the nutritious semen. Compared to their closest relatives, Neotrogla species can hold twice as much semen within their sperm storage organs. Scientists have speculated that a valve-like structure within this organ enables this extra storage by allowing the female to redirect semen into two separate chambers. But the organ’s small size has made it difficult to determine its inner workings, and scientists have yet to discover a switching valve that serves such a purpose within the animal kingdom. Yoshizawa et al. examined three Neotrogla species using advanced imaging technology and detected the first example of a biological switching valve. Neotrogla females can control this valve, switching the flow of semen between two slots. In this way, females can store two batches of semen at once. Seemingly exploiting this adaptation, the females’ spiky penises help them restrain males until they have received multiple semen injections. Yoshizawa et al. therefore suggest the emergence of this valve within the sperm storage organ may have promoted the evolution of the female penis. Along with giving insight into the lives of cave insects, these findings are of interest to engineers, who face challenges when constructing valves on a microscopic scale. The unique switching valve of female Neotrogla may one day inspire new man-made machinery that could advance a range of industries.

The elytral base sclerites (= sclerites located at the articular region between the forewing and thorax in Coleoptera) of selected taxa were examined and homologized. Although the elytral base sclerites are highly modified compared to the wing base sclerites of the other neopterans, they can be homologized by using the conservative wing flapping and folding lines as landmarks. A reduction of the first axillary sclerite was identified as a general trend of the elytral base sclerites, although the sclerite usually has a very important function to mediate flight power from the notum to the wing. This result indicates that the functional constraint against the basal sclerites is relaxed because of the lack of an ability to produce flight power by elytra. In contrast, the elytral folding system formed by the basal sclerites is well retained, which probably occurs because proper wing folding is a key for the shelter function of the elytra. The elytral base sclerites apparently contain more homoplasies than the serially homologous hindwing base sclerites of Coleoptera, which suggests that the structure is less useful for higher-level systematics. However, the faster evolutionary rate of the elytral base sclerites suggests there is potential for studying the lower-level phylogeny of Coleoptera.