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Although organ systems evolve in response to many intrinsic and extrinsic factors, frequently one factor has a dominating influence. For example, mouthpart shape and mechanics are thought to correlate strongly with aspects of the diet. Within insects, this paradigm of a shape-diet connection is advocated for decades but the relationship has so far never been quantified and is mostly based on qualitative observations. Orthoptera (grasshoppers, crickets, and allies) are a prominent case, for which mandible shape and dietary preference are thought to correlate strongly and even lead to predictions of feeding preferences. Here, we analysed mandible shape, biting efficiency, and their potential correlation with dietary categories in a phylogenetic framework for a broad sampling of several hundred extant Orthoptera covering nearly all families. The mandibular mechanical advantage was used as a descriptor of gnathal edge shape and bite force transmission efficiency. We aimed to understand how mandible shape is linked to biting efficiency and diet, and how these traits are influenced by phylogeny and allometry. The investigation reveals that feeding ecology is not the unequivocal predictor of mandible shape that it was assumed to be. There is a strong phylogenetic signal suggesting that phylogenetic history does have a much more prevalent influence on gnathal edge shape and distal mechanical advantage, than, e.g., feeding guilds or the efficiency of the force transmission to the food. Being ancestrally phytophagous, Orthoptera evolved in an environment with abundant food sources so that selective pressures leading to more specialized mouthpart shapes and force transmission efficiencies were low.

For over a century, the metapleural gland, an exocrine gland above the hind coxa, has been thought to be a unique structure for ants (Hymenoptera: Formicidae), and regarded as a catalyst for the ecological and evolutionary success of the family. This gland is one of the most researched exocrine glands in arthropods and its anatomy, ultrastructure, and chemistry are well documented. Herein, we describe an exocrine gland from the proctotrupoid wasp Pelecinus polyturator (Hymenoptera: Pelecinidae) with a similar position, structure, and chemistry to the ant metapleural gland: it is located just above the hind coxa, corresponds to an externally concave and fenestrated atrium, is composed of class 3 gland cells, and its extract contains relatively strong acids. We discover that the pelecinid gland is associated with the dilator muscle of the first abdominal spiracle, a trait that is shared with ants, but remained overlooked, possibly due to its small diameter, or obfuscation by the extensive metapleural gland. We also provide a biomechanical argument for passive emptying of the gland in both taxa. Pelecinids and ants with metapleural glands share a close association with soil. The pelecinid metapleural gland might therefore also have an antiseptic function as suggested for ants. We examined 44 other Hymenoptera families and found no glands associated with the oclusor apodeme or any signs of external modification. Our results strongly indicate that this complex trait (anatomical & chemical) evolved independently in ants and pelecinid wasps providing an exceptional system to better understand exocrine gland evolution in Hymenoptera.

In recent years, the study of insect morphology has benefited greatly from the emergence of new digital imaging and analysis technologies such as X-ray micro-computed tomography (μ-CT), digital 3D reconstruction, and animation. Through interactive gaming and virtual reality, the external morphology of insects can be studied by a broad audience of both entomologists and non-specialists. EntomonVR is a serious game designed to investigate the external morphology of insects with adequate quality for the virtual reality platform. We discuss the advantages of virtual reality, introduce the EntomonVR new educational game, and conclude about future perspectives, validations, and cost-effectiveness. For game assessment, we have tested this game on 25 participants with an entomological background and improved the game based on their feedback. This study demonstrates the efficacy of virtual reality technology for an experimental learning environment in teaching the morphology of insects and the crucial needs for advancing an efficient and interactive educational program.

Recent studies of insect anatomy evince a trend towards a comprehensive and integrative investigation of individual traits and their evolutionary relationships. The abdomen of ants, however, remains critically understudied. To address this shortcoming, we describe the abdominal anatomy of Amblyopone australis Erichson, using a multimodal approach combining manual dissection, histology, and microcomputed tomography. We focus on skeletomusculature, but additionally describe the metapleural and metasomal exocrine glands, and the morphology of the circulatory, digestive, reproductive, and nervous systems. We describe the muscles of the dorsal vessel and the ducts of the venom and Dufour's gland, and characterize the visceral anal musculature. Through comparison with other major ant lineages, apoid wasps, and other hymenopteran outgroups, we provide a first approximation of the complete abdominal skeletomuscular groundplan in Formicidae, with a nomenclatural schema generally applicable to the hexapod abdomen. All skeletal muscles were identifiable with their homologs, while we observe potential apomorphies in the pregenital skeleton and the sting musculature. Specifically, we propose the eighth coxocoxal muscle as an ant synapomorphy; we consider possible transformation series contributing to the distribution of states of the sternal apodemes in ants, Hymenoptera, and Hexapoda; and we address the possibly synapomorphic loss of the seventh sternal-eighth gonapophyseal muscles in the vespiform Aculeata. We homologize the ovipositor muscles across Hymenoptera, and summarize demonstrated and hypothetical muscle functions across the abdomen. We also give a new interpretation of the proximal processes of gonapophyses VIII and the ventromedial processes of gonocoxites IX, and make nomenclatural suggestions in the context of evolutionary anatomy and ontology. Finally, we discuss the utility of techniques applied and emphasize the value of primary anatomical research. ispartof: JOURNAL OF MORPHOLOGY vol:283 issue:6 pages:693-770 ispartof: location:United States status: published

Fossils provide unique opportunity to understand the tempo and mode of evolution and are essential for modeling the history of lineage diversification. Here, we interrogate the Mesozoic fossil record of the Aculeata, with emphasis on the ants (Formicidae), and conduct an extended series of ancestral state estimation exercises on distributions of tip-dated combined-evidence phylogenies. We developed and illustrated from ground-up a series of 576 morphological characters which we scored for 144 extant and 431 fossil taxa, including all families of Aculeata, Trigonaloidea, Evanioidea, and †Ephialtitoidea. We used average posterior probability support to guide composition of a target matrix of 303 taxa, for which we integrated strongly filtered ultraconserved element (UCE) data for 115 living species. We also implemented reversible jump MCMC (rjMCMC) and hidden state methods to model complex behavioral characters to test hypotheses about the pathway to obligate eusociality. In addition to revising the higher classification of all sampled groups to family or subfamily level using estimated character polarities to diagnose nodes across the phylogeny, we find that the mid-Cretaceous genera †Camelomeciaand †Camelospheciaform a clade which is robustly supported as sister to all living and fossil Formicidae. For this reason, we name this extinct clade as †@@@idaefam. nov.and provide a definition for the expanded Formicoidea. Based on our results, we recognize three major phases in the early evolution of the ants: (1) origin of Formicoidea as ground-adapted huntresses during the Late Jurassic in the “stinging aggressor” guild (Aculeata) among various lineages of “sneaking parasitoids” (non-aculeate Vespina); (2) the first formicoid radiation during the Early Cretaceous, by the end of which all major extant linages originated; and (3) turnover of the Formicoidea at the end-Cretaceous leading to the second formicoid radiation. We conclude with a concentrated series of considerations for future directions of study with this dataset and beyond.

One key event in insect evolution was the development of mandibles with two joints, which allowed powerful biting but restricted their movement to a single degree of freedom. These mandibles define the Dicondylia, which constitute over 99% of all extant insect species. It was common doctrine that the dicondylic articulation of chewing mandibles remained unaltered for more than 400 million years. We report highly modified mandibles overcoming the restrictions of a single degree of freedom and hypothesize their major role in insect diversification. These mandibles are defining features of parasitoid chalcid wasps, one of the most species-rich lineages of insects. The shift from powerful chewing to precise cutting likely facilitated adaptations to parasitize hosts hidden in hard substrates, which pose challenges to the emerging wasps. We reveal a crucial step in insect evolution and highlight the importance of comprehensive studies even of putatively well-known systems.

The extraordinary success of social insects is partially based on division of labour, i.e. individuals exclusively or preferentially perform specific tasks. Task preference may correlate with morphological adaptations so implying task specialization, but the extent of such specialization can be difficult to determine. Here, we demonstrate how the physical foundation of some tasks can be leveraged to quantitatively link morphology and performance. We study the allometry of bite force capacity in Atta vollenweideri leaf-cutter ants, polymorphic insects in which the mechanical processing of plant material is a key aspect of the behavioural portfolio. Through a morphometric analysis of tomographic scans, we show that the bite force capacity of the heaviest colony workers is twice as large as predicted by isometry. This disproportionate ‘boost’ is predominantly achieved through increased investment in muscle volume; geometrical parameters such as mechanical advantage, fibre length or pennation angle are likely constrained by the need to maintain a constant mandibular opening range. We analyse this preference for an increase in size-specific muscle volume and the adaptations in internal and external head anatomy required to accommodate it with simple geometric and physical models, so providing a quantitative understanding of the functional anatomy of the musculoskeletal bite apparatus in insects.

The extraordinary success of social insects is partially based on ‘division of labour’, i. e. individuals exclusively or preferentially perform specific tasks. Task-preference may correlate with morphological adaptations so implying task-specialisation, but the extent of such specialisation can be difficult to determine. Here, we demonstrate how the physical foundation of some tasks can be leveraged to quantitatively link morphology and performance. We study the allometry of bite force capacity inAtta vollenweiderileaf-cutter ants, polymorphic insects in which the mechanical processing of plant material is a key aspect of the behavioural portfolio. Through a morphometric analysis of tomographic scans, we show that the bite force capacity of the heaviest colony workers is twice as large as predicted by isometry. This disproportionate ‘boost’ is predominantly achieved through increased investment in muscle volume; geometrical parameters such as mechanical advantage, fibre length or pennation angle are likely constrained by the need to maintain a constant mandibular opening range. We analyse this preference for an increase in size-specific muscle volume and the adaptations in internal and external head anatomy required to accommodate it with simple geometric and physical models, so providing a quantitative understanding of the functional anatomy of the musculoskeletal bite apparatus in insects.

Direct fossil evidence for Mesozoic flower pollination is scarce. Umenocoleoid micro-cockroaches Lepidopterix vegrandis gen. et sp. n. (Lebanese amber) and Antophiloblatta hispida gen. et sp. n. (Myanmar amber) possess size, form, complex coloration pattern and surface structure, cryptic with potentially entomophilous angiosperms Tropidogyne pentaptera Poinar, 2017 and Antiquifloris latifibris Poinar et Buckley, 2016 flower petals and sepals. Putative pollen grains attached to the latter adult indicates pollination, while reduced mouthparts suggest fluid nectar feeding. Spongistoma angusta gen. et sp. n. (Myanmar amber) has a narrow body and mandibles nearly entirely reduced with a unique “proboscis” forming sponging/ sucking mouthparts. In addition putative Classopolis Pflug, 1953 gymnosperm pollen is attached to adults and immature individuals of Vzrkadlenie miso gen. et sp. n. (Myanmar amber). Together with possible angiosperm pollination by Formicamendax vrsanskyi Hinkelman, 2019 and cycas pollination by immature individuals of alienopterid larvae, the evidence for early cockroach pollination is now substantial. Additionally unique is the forewing surface of L. vegrandis with photonic crystal structures within the scales.

Proceedings of the Royal Society of London / B 288, 20210616 (2021). doi:10.1098/rspb.2021.0616, Most animals undergo ecological niche shifts between distinct life phases, but such shifts can result in adaptive conflicts of phenotypic traits. Metamorphosis can reduce these conflicts by breaking up trait correlations, allowing each life phase to independently adapt to its ecological niche. This process is called adaptive decoupling. It is, however, yet unknown to what extent adaptive decoupling is realized on a macroevolutionary scale in hemimetabolous insects and if the degree of adaptive decoupling is correlated with the strength of ontogenetic niche shifts. It is also unclear whether the degree of adaptive decoupling is correlated with phenotypic disparity. Here, we quantify nymphal and adult trait correlations in 219 species across the whole phylogeny of earwigs and stoneflies to test whether juvenile and adult traits are decoupled from each other. We demonstrate that adult head morphology is largely driven by nymphal ecology, and that adult head shape disparity has increased with stronger ontogenetic niche shifts in some stonefly lineages. Our findings implicate that the hemimetabolan metamorphosis in earwigs and stoneflies does not allow for high degrees of adaptive decoupling, and that high phenotypic disparity can even be realized when the evolution of distinct life phases is coupled., Published by Royal Soc. of London, London

Spiders use their inner body fluid (“blood” or hemolymph) to drive hydraulic extension of their legs. In hydraulic systems, performance is highly dependent on the working fluid, which needs to be chosen according to the required operating speed and pressure. Here, we provide new insights into the fluid mechanics of spider locomotion. We present the three-dimensional structure of one of the crucial joints in spider hydraulic actuation, elucidate the fluid flow inside the spider leg, and quantify the rheological properties of hemolymph under physiological conditions. We observe that hemolymph behaves as a shear-thinning non-Newtonian fluid with a fluid behavior indexn= 0.5, unlike water (n= 1.0)., Soft Matter, 17 (22), ISSN:1744-683X, ISSN:1744-6848

Palaeontologia electronica 24(1), a07 (2021). doi:10.26879/1129, We report a hitherto unprecedented diversity of fly larvae (Diptera) from Eocene Baltic amber and the use of these to address palaeo-ecosystem functions and processes in the surrounding forests. Fly larvae have been considered exceptionally rare by the research community and have, like most insect larvae, been deemed of limited utility owing to challenges in identification. Herein, however, using synchrotron-x-ray radiation CT (SR-µCT) allowed us to detect and identify dozens of fly larvae from Baltic amber, and to infer their ecological interactions. One particular piece of amber contains 56 fly larvae and apparent mammalian feces. This fossil is of great interest for our understanding of carbon cycling in the Eocene forest. The occurrence of such a large number of fly larvae on the fecal remains indicates an important role of flies in recycling organic matter in the Eocene forest, much as some larvae do today. Analysis of the fly palaeo-communities also allowed us to hypothesize a mechanism by which massive, geologically relevant deposits of amber were formed in the Baltic region. Scanning allowed us to identify seven larvae closely related to the extant Syrphidae, whose larvae inhabit nests of eusocial Hymenoptera, or, sometimes, flows of sap dripping from trees damaged by other burrowing insect larvae., Published by Coquina Press, [S.l.]

We present Biomedisa, a free and easy-to-use open-source online platform developed for semi-automatic segmentation of large volumetric images. The segmentation is based on a smart interpolation of sparsely pre-segmented slices taking into account the complete underlying image data. Biomedisa is particularly valuable when little a priori knowledge is available, e.g. for the dense annotation of the training data for a deep neural network. The platform is accessible through a web browser and requires no complex and tedious configuration of software and model parameters, thus addressing the needs of scientists without substantial computational expertise. We demonstrate that Biomedisa can drastically reduce both the time and human effort required to segment large images. It achieves a significant improvement over the conventional approach of densely pre-segmented slices with subsequent morphological interpolation as well as compared to segmentation tools that also consider the underlying image data. Biomedisa can be used for different 3D imaging modalities and various biomedical applications., Manual segmentation of biological images is a time-consuming task. Here the authors present Biomedisa, an open-source online platform for segmentation of large volumetric images starting from sparsely presegmented slices.

Thickened femora of insects are correlated to enlarged muscle masses and serve two basic purposes: jumping/kicking and grasping/holding. Modifications on the ventral femoral wall and the tibial flexor tendon that are possibly involved in catch mechanisms have been described in multiple insect taxa with both jumping and grasping legs. Our comparative study aims to explore the functional and structural similarities of these modification in jumping and grasping leg types from Coleoptera, Hymenoptera, Diptera, and Orthoptera with the combination of cutting edge, noninvasive imaging methods, and classical dissections techniques. Our data indicate that locking mechanisms are present in the jumping and grasping legs of insects. We describe three femoro-tibial lock types based on the location of the interacting sclerites relative to the site of origin of the tibial flexor tendon. All of the three types can be found in jumping insect legs, whereas only one type is present in grasping legs. The locking mechanism might aid in keeping the femoro-tibial joint in a flexed position for an extended period of time. Our data indicate that morphologically similar modifications in the femoro-tibial joint are involved in energy-saving mechanisms both in jumping and grasping legs in insects.

2.AbstractSuction organs provide powerful yet dynamic attachments for many aquatic animals, including octopus, squid, remora, and clingfish. While the functional morphology of suction organs from various cephalopods and fishes has been investigated in detail, there are only few studies on such attachment devices in insects. Here we characterise the morphology, ultrastructure, andin vivomovements of the suction attachment organs of net-winged midge larvae (genusLiponeura) – aquatic insects that live on rocks in rapid alpine waterways where flow rates can reach 3 m s-1– using scanning electron microscopy, laser confocal scanning microscopy, and X-ray computed micro-tomography (micro-CT). We identified structural adaptations important for the function of the suction attachment organs fromL. cinerascensandL. cordata. First, a dense array of spine-like microtrichia covering each suction disc comes into contact with the substrate upon attachment. Similar hairy structures have been found on the contact zones of suction organs from octopus, clingfish, and remora fish. These structures are thought to contribute to the seal and to provide increased shear force resistance in high-drag environments. Second, specialised rim microtrichia at the suction disc periphery form a continuous ring in close contact with a surface and may serve as a seal on a variety of surfaces. Third, a V-shaped cut on the suction disc (the V-notch) is actively peeled open via two cuticular apodemes inserting into its flanks. The apodemes are attached to dedicated V-notch opening muscles, thereby providing a unique detachment mechanism. The complex cuticular design of the suction organs, along with specialised muscles that attach to them, allows blepharicerid larvae to generate powerful attachments which can withstand strong hydrodynamic forces and quickly detach for locomotion. Our findings could be applied to bio-inspired attachment devices that perform well on a wide range of surfaces.

BackgroundSuction organs provide powerful yet dynamic attachments for many aquatic animals, including octopus, squid, remora, and clingfish. While the functional morphology of suction organs from some cephalopods and fishes has been investigated in detail, there are only few studies on such attachment devices in insects. Here we characterise the morphology and ultrastructure of the suction attachment organs of net-winged midge larvae (genusLiponeura; Diptera: Blephariceridae) – aquatic insects that live on rocks in rapid alpine waterways where flow speeds can reach 3 m s− 1– using scanning electron microscopy, confocal laser scanning microscopy, and X-ray computed micro-tomography (micro-CT). Furthermore, we study the function of these organs in vivo using interference reflection microscopy.ResultsWe identified structural adaptations important for the function of the suction attachment organs inL. cinerascensandL. cordata. First, a dense array of spine-like microtrichia covering each suction disc comes into contact with the substrate upon attachment, analogous to hairy structures on suction organs from octopus, clingfish, and remora fish. These spine-like microtrichia may contribute to the seal and provide increased shear force resistance in high-drag environments. Second, specialised rim microtrichia at the suction disc periphery were found to form a continuous ring in close contact and may serve as a seal on a variety of surfaces. Third, a V-shaped cut on the suction disc (“V-notch“) is actively opened via two cuticular apodemes inserting on its flanks. The apodemes are attached to dedicated V-notch opening muscles, thereby providing a unique detachment mechanism. The complex cuticular design of the suction organs, along with specialised muscles that attach to them, allows blepharicerid larvae to generate powerful attachments which can withstand strong hydrodynamic forces and quickly detach for locomotion.ConclusionThe suction organs fromLiponeuraare underwater attachment devices specialised for resisting extremely fast flows. Structural adaptations from these suction organs could translate into future bioinspired attachment systems that perform well on a wide range of surfaces.

Ceraphronoids are some of the most commonly collected hymenopterans, yet they remain rare in the fossil record.Conostigmus talamasiMikó and Trietsch, sp. nov. from Baltic amber represents an intermediate form between the type genus,Megaspilus, and one of the most species-rich megaspilid genera,Conostigmus. We describe the new species using 3D data collected with synchrotron-based micro-CT equipment. This non-invasive technique allows for quick data collection in unusually high resolution, revealing morphological traits that are otherwise obscured by the amber. In describing this new species, we revise the diagnostic characters for Ceraphronoidea and discuss possible reasons why minute wasps with a pterostigma are often misidentified as ceraphronoids. Based on the lack of ceraphronoid characteristics, we removeDendrocerus dubitatusBrues, 1937, Stigmaphronidae, and Radiophronidae from Ceraphronoidea and consider them asincertae sedis. We also provide some guidance for their future classification.

Modern applications for analysing 2D/3D data require complex visual output features which are often based on the multi-platform OpenGL® API for rendering vector graphics. Instead of providing classical workstations, the provision of powerful virtual machines (VMs) with GPU support in a scientific cloud with direct access to high performance storage is an efficient and cost effective solution. However, the automatic deployment, operation and remote access of OpenGL® API-capable VMs with professional visualization applications is a non-trivial task. In this chapter the authors demonstrate the concept of such a flexible cloud-like analysis infrastructure within the framework of the project ASTOR. The authors present an Analysis-as-a-Service (AaaS) approach based on VMware™-ESX for on demand allocation of VMs with dedicated GPU cores and up to 256 GByte RAM per machine.

Developments in X-Ray Tomography XI : [Proceedings] - SPIE, 2017. - ISBN 9781510612396 - doi:10.1117/12.2275121 Developments in X-Ray Tomography XI, San Diego, United States, 6 Aug 2017 - 10 Aug 2017; Proceedings of SPIE 10391, 103910P (2017). doi:10.1117/12.2275959, Beamtime and resulting SRμCT data are a valuable resource for researchers of a broad scientific community in life sciences. Most research groups, however, are only interested in a specific organ and use only a fraction of their data. The rest of the data usually remains untapped. By using a new collaborative approach, the NOVA project (Network for Online Visualization and synergistic Analysis of tomographic data) aims to demonstrate, that more efficient use of the valuable beam time is possible by coordinated research on different organ systems. The biological partners in the project cover different scientific aspects and thus serve as model community for the collaborative approach. As proof of principle, different aspects of insect head morphology will be investigated (e.g., biomechanics of the mouthparts, and neurobiology with the topology of sensory areas). This effort is accomplished by development of advanced analysis tools for the ever-increasing quantity of tomographic datasets. In the preceding project ASTOR, we already successfully demonstrated considerable progress in semi-automatic segmentation and classification of internal structures. Further improvement of these methods is essential for an efficient use of beam time and will be refined in the current NOVAproject. Significant enhancements are also planned at PETRA III beamline p05 to provide all possible contrast modalities in x-ray imaging optimized to biological samples, on the reconstruction algorithms, and the tools for subsequent analyses and management of the data. All improvements made on key technologies within this project will in the long-term be equally beneficial for all users of tomography instrumentations., Published by SPIE, Bellingham, Wash.

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New imaging stations aim for high spatial and temporal resolution and are characterized by ever increasing sampling rates and demanding data processing workflows. Key to successful imaging experiments is to open up high-performance computing resources. This includes carefully selected components for computing hardware and development of advanced imaging algorithms optimized for efficient use of parallel processor architectures. We present the novel UFO computing platform for online data processing for imaging experiments and image-based feedback. The platform handles the full data life cycle from the X-ray detector to long-term data archives. Core components of this system are an FPGA platform for ultra-fast data acquisition, the GPU-based UFO image processing framework, and the fast control system “Concert”. Reconstruction algorithms implemented in the UFO framework are optimized for the latest GPU architectures and provide a reconstruction throughput in the GB/s-range. The control system “Concert” integrates high-speed computing nodes and fast beamline devices and thus enables image-based control loops and advanced workflow automation for efficient beam time usage. Low latencies are ensured by direct communication between FPGA and GPUs using AMDs DirectGMA technology. Time resolved tomography is supported by cutting edge regularization methods for high quality reconstructions with a reduced number of projections. The new infrastructure at ANKA has dramatically accelerated tomography from hours to second and resulted in new application fields, like high-throughput tomography, pump-probe radiography and stroboscopic tomography. Ultra-fast X-ray cine-tomography for the first time allows one to observe internal dynamics of moving millimeter-sized objects in real-time.

We analyze theoretically and investigate experimentally the transfer of phase to intensity power spectra of spatial frequencies through free-space Fresnel diffraction. Depending on $\ensuremath{\lambda}z$ (where $\ensuremath{\lambda}$ is the wavelength and $z$ is the free-space propagation distance) and the phase-modulation strength $S$, we demonstrate that for multiscale and broad phase spectra critical behavior transmutes a quasilinear to a nonlinear diffractogram except for low frequencies. On the contrary, a single-scale and broad phase spectrum induces a critical transition in the diffractogram at low frequencies. In both cases, identifying critical behavior encoded in the intensity power spectra is of fundamental interest because it exhibits the limits of perturbative power counting but also guides resolution and contrast optimization in propagation-based, single-distance, phase-contrast imaging, given certain dose and coherence constraints.

With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These “missing links” fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.

Two genera of extinct weevils, Sayrevilleus Gratshev & Zherikhin from Cretaceous New Jersey amber and Baltocar Kuschel from Eocene Baltic amber, are recognized as close relatives based on similarities revealed by the use of synchrotron tomography and the availability of new amber inclusions. The subfamily Sayrevilleinae Legalov stat. nov. is characterized by possessing mandibles with an external cutting edge and an inner blunt edge. The subfamily is placed in the family Attelabidae (s.l.), although some characters also suggest a possible relationship with the ‘higher weevils’ comprising Caridae, Brentidae, and Curculionidae. Sayrevilleus is transferred from the tribe Auletini of Rhynchitinae to Sayrevilleinae, and Sayrevilleus grimaldii Gratshev & Zherikhin is redescribed. Baltocar Kuschel is transferred from Caridae to Sayrevilleinae and revised, its type species, Baltocar succinicus (Voss), is redescribed and three new species, Baltocar groehni Riedel sp. nov., Baltocar hoffeinsorum Riedel sp. nov., and Baltocar subnudus Riedel sp. nov. are described based on eight well-preserved inclusions. The genera Orapauletes Legalov and Zherichiniletes Legalov previously assigned to Sayrevilleini are regarded as Curculionoidea incertae sedis. The Sayrevilleinae were distributed over areas of North America and Europe at least since the Late Cretaceous (c. 90 Mya) and were probably relatively diverse until the Eocene (c. 44 Mya). It is speculated that they became extinct through competition with Curculionidae, which used a similar oviposition strategy. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165, 773–794.