Revisiting the four Hexapoda classes: Protura as the sister group to all other hexapods.

Insects represent the most diverse animal group, yet previous phylogenetic analyses based on morphological and molecular data have failed to agree on the evolutionary relationships of early insects and their six-legged relatives (together constituting the clade Hexapoda). In particular, the phylogenetic positions of the three early-diverging hexapod lineages-the coneheads (Protura), springtails (Collembola), and two-pronged bristletails (Diplura)-have been debated for over a century, with alternative topologies implying drastically different scenarios of the evolution of the insect body plan and hexapod terrestrialization. We addressed this issue by sampling all hexapod orders and experimenting with a broad range of across-site compositional heterogeneous models designed to tackle ancient divergences. Our analyses support Protura as the earliest-diverging hexapod lineage ("Protura-sister") and Collembola as a sister group to Diplura, a clade corresponding to the original composition of Entognatha, and characterized by the shared possession of internal muscles in the antennal flagellum. The previously recognized 'Elliplura' hypothesis is recovered only under the site-homogeneous substitution models with partial supermatrices. Our cross-validation analysis shows that the site-heterogeneous CAT-GTR model, which recovers "Protura-sister," fits significantly better than homogeneous models. Furthermore, the morphologically unusual Protura are also supported as the earliest-diverging hexapod lineage by other lines of evidence, such as mitogenomes, comparative embryology, and sperm morphology, which produced results similar to those in this study. Our backbone phylogeny of hexapods will facilitate the exploration of the underpinnings of hexapod terrestrialization and megadiversity.

WingAnalogy: a computer vision-based tool for automated insect wing asymmetry and morphometry analysis.

WingAnalogy is a computer tool for automated insect wing morphology and asymmetry analysis. It facilitates project management, enabling users to import pairs of wing images obtained from individual insects, such as left and right, fore- and hindwings. WingAnalogy employs image processing and computer vision to segment wing structures and extract cell boundaries, and junctions. It quantifies essential metrics encompassing cell and wing characteristics, including area, length, width, circularity, and centroid positions. It enables users to scale and superimpose wing images utilizing Particle Swarm Optimization (PSO). WingAnalogy computes regression, Normalized Root Mean Square Error (NRMSE), various cell-based parameters, and distances between cell centroids and junctions. The software generates informative visualizations, aiding researchers in comprehending and interpreting asymmetry patterns. WingAnalogy allows for dividing wings into up to five distinct wing cell sets, facilitating localized comparisons. The software excels in report generation, providing detailed asymmetry measurements in PDF, CSV, and TXT formats.

Early Pennsylvanian Lagerstätte reveals a diverse ecosystem on a subhumid, alluvial fan.

Much of what we know about terrestrial life during the Carboniferous Period comes from Middle Pennsylvanian (~315-307 Mya) Coal Measures deposited in low-lying wetland environments1-5. We know relatively little about terrestrial ecosystems from the Early Pennsylvanian, which was a critical interval for the diversification of insects, arachnids, tetrapods, and seed plants6-10. Here we report a diverse Early Pennsylvanian trace and body fossil Lagerstätte (~320-318 Mya) from the Wamsutta Formation of eastern North America, distinct from coal-bearing deposits, preserved in clastic substrates within basin margin conglomerates. The exceptionally preserved trace fossils and body fossils document a range of vertebrates, invertebrates and plant taxa (n = 131), with 83 distinct foliage morphotypes. Plant-insect interactions include what may be the earliest evidence of insect oviposition. This site expands our knowledge of early terrestrial ecosystems and organismal interactions and provides ground truth for future phylogenetic reconstructions of key plant, arthropod, and vertebrate groups.

The structure of a gilled stonefly larva from the mid-Cretaceous Kachin amber.

Perlidae stands as the most diverse family within Plecoptera, with evidence suggesting possible adaptation to warmer aquatic environments. Tracheal gills are hypothesized to have played a pivotal role in this radiation process. This study presents the description of a fossilized stonefly larva with gills, preserved as a fresh exuvia in mid-Cretaceous Kachin amber from northern Myanmar. The larva was classified within the family Perlidae based on distinctive morphological traits, including toothed lacinia and sharp-cusped mandibles, slender palps, glossae shorter than rounded paraglossae, and highly branched gills on the sides and ventral surface of thoracic segments. Additionally, the presence of a transverse, sparse, and irregular setal row on the occiput further indicates classification within the subfamily Acroneuriinae. Notably, the fossilized larva displays striking similarities in gill morphology and distribution to certain extant members within Perlidae, suggesting that these gill structures have an advantage in various aquatic habitats.

The contribution of the Middle Triassic fossil assemblage of Monte San Giorgio to insect evolution.

The Triassic represents a critical period for understanding the turnover of insect fauna from the Paleozoic to the Mesozoic following the end-Permian mass extinctions (EPME); however, fossil deposits from the Early-Middle Triassic are scarce. The exceptionally preserved 239 million-year-old fossil insect fauna recorded at Monte San Giorgio (Switzerland), including 248 fossils representing 15 major insect clades is presented here. Besides the exceptional features, including their small size and excellent preservation, the fossils have importance in the evolutionary history of the group. The taxonomic and ecological diversity recovered, including both freshwater (dragonflies and caddisflies) and terrestrial taxa (true bugs and wasps), demonstrates that complex environments sustained a paleocommunity dominated by monurans (thought not to have survived the EPME), midges, and beetles. Interestingly, a blattodean-like fossil bearing an external ootheca was also found, important for understanding Paleozoic roachoids to extant cockroaches' transition and the evolution of maternal brood care. Moreover, the youngest and first complete specimen of †Permithonidae and the oldest sawfly fossils were discovered. Finally, round-shaped bodies, compatible with seminal capsules or lycophyte spores, were found on the abdomens of several midge-like individuals. If these are spores, non-seed-bearing plants could have been the first entomophilous plants rather than gymnosperms, as recently supposed. Altogether, these fossils contribute substantially to understanding insect evolution and Paleozoic-Mesozoic faunal turnover.

Mesozoic larva in amber reveals the venom delivery system and the palaeobiology of an ancient lineage of venomous insects (Neuroptera).

The larvae of Neuroptera are predators that feed by injecting bioactive compounds into their prey and then suctioning the fluids through modified mouthparts. We explore the evolutionary history of this feeding structure through the examination of a new fossil larva preserved in Late Cretaceous Kachin amber, which we describe as new genus and species, Electroxipheus veneficus gen et sp. nov. X-ray phase-contrast microtomography enabled us to study the anatomy of the larva in 3D, including the structure of the mouthparts and that of the venom delivery system. The specimen exhibited a unique combination of morphological traits not found in any known fossil or extant lacewing, including an unusual structure of the antenna. Phylogenetic analyses, incorporating a selection of living and fossil larval Neuroptera and enforcing maximum parsimony and Bayesian inference, identified the larva as belonging to the stem group Mantispoidea. The larva shows that the anatomy of the feeding and venom-delivery apparatus has remained unchanged in Neuroptera from the Cretaceous to the present. The morphology of the specimen suggests that it was an active predator, in contrast with the scarcely mobile, specialized relatives, like mantispids and berothids.

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

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

Comparative morphology of male genitalia in antlions (Insecta, Neuroptera, Myrmeleontidae), with emphasis on owlflies (Ascalaphinae) and a possible structural evolutionary scenario.

Male genitalia morphology in Myrmeleontidae has traditionally been insufficiently studied, although it has received increased attention for its diagnostic value in recent times. A neutral terminology has generally been applied in standard taxonomic practice, yet knowledge of an equivalent and stable terminology across taxa based on comparative morphology has been missing. Herein a detailed comparative morphology study with examples from most tribes within Myrmeleontidae, including owlflies (Ascalaphinae), attempts to relate external and internal genital structures based on a proposed groundplan for Neuroptera and Myrmeleontidae. We contend that a groundplan based on 10 abdominal segments, plus vestigial structures from an 11th segment, coherently depicts structural components across myrmeleontid taxa. A gonarcus, an element of Neuropterida amply referred in Neuroptera, is supported to represent the pair of abdominal appendages of segment X medially fused, with gonocoxite and gonostylus components. In most myrmeleontid taxa, basal (gonocoxites) and distal (gonostyli) components separate, with gonostyli positioned posteriorly with respect to gonocoxites, still united with translucent, lightly sclerotized tissue, forming a more or less conical structure, a proposed synapomorphy for the family. Ninth gonostyli are generally reduced (pulvini) and have migrated close to the base of gonarcus (10th gonocoxites). A pelta, also a potential synapomorphy for Myrmeleontidae, derives from paired setose surfaces of the 10th gonostyli, medially positioned (still evident in Bubopsis). Three structural types of gonarcus are diagnosed for illustrative purposes, as they may represent convergent constructs.

Automated identification of aquatic insects: A case study using deep learning and computer vision techniques.

Deep learning techniques have recently found application in biodiversity research. Mayflies (Ephemeroptera), stoneflies (Plecoptera) and caddisflies (Trichoptera), often abbreviated as EPT, are frequently used for freshwater biomonitoring due to their large numbers and sensitivity to environmental changes. However, the morphological identification of EPT species is a challenging but fundamental task. Morphological identification of these freshwater insects is therefore not only extremely time-consuming and costly, but also often leads to misjudgments or generates datasets with low taxonomic resolution. Here, we investigated the application of deep learning to increase the efficiency and taxonomic resolution of biomonitoring programs. Our database contains 90 EPT taxa (genus or species level), with the number of images per category ranging from 21 to 300 (16,650 in total). Upon completion of training, a CNN (Convolutional Neural Network) model was created, capable of automatically classifying these taxa into their appropriate taxonomic categories with an accuracy of 98.7 %. Our model achieved a perfect classification rate of 100 % for 68 of the taxa in our dataset. We achieved noteworthy classification accuracy with morphologically closely related taxa within the training data (e.g., species of the genus Baetis, Hydropsyche, Perla). Gradient-weighted Class Activation Mapping (Grad-CAM) visualized the morphological features responsible for the classification of the treated species in the CNN models. Within Ephemeroptera, the head was the most important feature, while the thorax and abdomen were equally important for the classification of Plecoptera taxa. For the order Trichoptera, the head and thorax were almost equally important. Our database is recognized as the most extensive aquatic insect database, notably distinguished by its wealth of included categories (taxa). Our approach can help solve long-standing challenges in biodiversity research and address pressing issues in monitoring programs by saving time in sample identification.

Worldwide revision of synanthropic silverfish (Insecta: Zygentoma: Lepismatidae) combining morphological and molecular data.

Synanthropic silverfish are the best-known and most widely distributed insects of the order Zygentoma. However, there is a great gap in the knowledge and confusion about the geographic distribution and the diagnostic characteristics that allow their identification. In this work, we provide an exhaustive and deep analysis of the most common 9 synanthropic silverfish of the world, combining previously published and newly derived morphological and molecular data. Updated descriptions of Ctenolepisma calvum (Ritter, 1910) and Ctenolepisma (Sceletolepisma) villosum (Fabricius, 1775) are included, and morphological remarks, illustrations, and photographs of the remaining synanthropic species are provided to clarify their diagnosis and differentiation among them and from other free-living species. In addition, Ctenolepisma targionii (Grassi and Rovelli, 1889) is synonymized with C. villosum. A molecular phylogeny is presented based on the COI sequences of all the synanthropic species deposited in BOLD and GenBank, with 15 new sequences provided by this study. This has allowed us to detect and correct a series of identification errors based on the lack of morphological knowledge of several species. Moreover, 2 different lineages of Ctenolepisma longicaudatumEscherich, 1905 have also been detected. To help future studies, we also provide a taxonomic interpretation guide for the most important diagnostic characters of the order Zygentoma, as well as an identification key for all the Synanthropic studied species. Finally, an approximation of the global distribution of synanthropic silverfish is discussed. Several new records indicate that the expansion of these species, generally associated with the transport of goods and people, is still far from over.

The structure of wing in the earliest Permopsocida.

Permopsocids are small acercarian insects with mouthparts specialized for sucking. They are closely related to Hemiptera and Thysanoptera. The earliest known representatives are from the Early Permian. Here evidence is presented that the Permopsocida occurred even earlier in Pennsylvanian (Moscovian) deposits in the Piesberg quarry near Osnabrück (Lower Saxony, Germany). This material is assigned to the Permian family Psocidiidae; Carbonopsocus mercuryi gen. et sp. nov., based on the wing venation diagnosed by the unique branching pattern of the main veins, the shape of the areola postica being longer than wide, the angular shape of the pterostigma, the ir crossvein directed proximally mid of pterostigma (apomorphy) and the vannus formed by the three veins of PCu, A1 and A2. The shape of the veins, with a Y-vein formed by the distal fusion of PCu with A1, could be a putative symplesiomorphy of the Psocodea with Permopsocida and Hemiptera. C. mercuryi gen. et sp. nov. is the first appearance date for Permopsocida and roots the Acercaria tree. In addition, another specimen of Dichentomum cf. arroyo (Psocidiidae) from Carrizo Arroyo is presented and figured, confirming the presence of the genus Dichentomum near the Carboniferous-Permian boundary and linking it to the Artinskian species from Elmo in Kansas, USA.

Insect Flight Energetics and the Evolution of Size, Form, and Function.

Flying insects vary greatly in body size and wing proportions, significantly impacting their flight energetics. Generally, the larger the insect, the slower its flight wingbeat frequency. However, variation in frequency is also explained by differences in wing proportions, where larger-winged insects tend to have lower frequencies. These associations affect the energy required for flight. The correlated evolution of flight form and function can be further defined using a lineage of closely related bee species varying in body mass. The decline in flight wingbeat frequency with increasing size is paralleled by the flight mass-specific metabolic rate. The specific scaling exponents observed can be predicted from the wing area allometry, where a greater increase (hyperallometry) leads to a more pronounced effect on flight energetics, and hypoallometry can lead to no change in frequency and metabolic rate across species. The metabolic properties of the flight muscles also vary with body mass and wing proportions, as observed from the activity of glycolytic enzymes and the phospholipid compositions of muscle tissue, connecting morphological differences with muscle metabolic properties. The evolutionary scaling observed across species is recapitulated within species. The static allometry observed within the bumblebee Bombus impatiens, where the wing area is proportional and isometric, affects wingbeat frequency and metabolic rate, which is predicted to decrease with an increase in size. Intraspecific variation in flight muscle tissue properties is also related to flight metabolic rate. The role of developmental processes and phenotypic plasticity in explaining intraspecific differences is central to our understanding of flight energetics. These studies provide a framework where static allometry observed within species gives rise to evolutionary allometry, connecting the evolution of size, form, and function associated with insect flight.

A new Drusinae (Trichoptera: Limnephilidae) species from the Zagros Mountains in the Kurdistan Province, Iran.

A new species of the Drusinae subfamily, Drusus chelchamaensis Ibrahimi & Mohammadi sp. nov., is described based on male genital morphology. The male of the new species is most similar to that of Drusus bayburtii akin 1983 (known from several places in Anatolia, Turkey), Drusus ketes Olh & Mey 2017 (in Olh et al. 2017) (known from the Hamedan Province, Iran), and Drusus kazanciae akin 1983 (known from the Hakkari Province, Turkey) and differs in the formation of superior appendages, intermediate appendages, inferior appendages, parameres, spinate area of segment VIII, and segment IX. The species is currently known only from Chel Chama Mountain in the Kurdistan Province of Iran and is the second known species of the Drusus caucasicus Species Complex from Iran. Chel Chama is part of the Zagros Mountains, which is rich in freshwater ecosystems, but currently with only a few reported species of caddisflies.

Description of the full-grown larva and barcode of Athripsodes taounate taounate Dakki & Malicky 1980 (Trichoptera: Leptoceridae), an Iberic-Maghrebian endemic.

In this paper we describe the main morphological characteristics that distinguish the full-grown larva of Athripsodes taounate taounate, an Iberic-Maghrebian endemic. The conspecificity of the larva and adult was confirmed by DNA analysis. Morphological features that discriminate it from the described Iberian-Maghrebian species of Athripsodes are given.

Morphology of the male reproductive system and spermatozoa of Smicridea (Rhyacophylax) iguazu Flint, 1983 (Trichoptera, Hydropsychidae).

The Trichoptera, holometabolous aquatic insects found worldwide except in Antarctica, exhibit a unique feature in their sperm, which are solely nucleated (eupyrene). Current knowledge on Trichoptera sperm is limited to Old World species. To enhance our understanding of their reproductive biology and contribute to systematic discussions, we describe the male reproductive system and spermatozoa of Smicridea (Rhyacophylax) iguazu Flint, 1983 (Hydropsychidae). This species lacks seminal vesicles, possesses piriform to oval-shaped testes with spermatozoa grouped in apical bundles and dense filamentous material filling other areas. The vasa deferentia are long and a pair of elongated accessory glands displays distinct proximal and distal regions. The relatively short (∼40 µm) spermatozoa are nucleated, aflagellated, and immobile. Further research could explore variations and assess the taxonomic utility of these features for genus identification within Hydropsychidae.

A new mayfly subfamily sheds light on the early evolution and Pangean origin of Baetiscidae (Insecta: Ephemeroptera).

The family Baetiscidae Edmunds & Traver, 1954 is recognisable among mayflies due to its bizarre larvae, equipped with a robust and spiked thoracic notal shield covering part of the abdomen up to sixth segment. Originally being described as extant species from the USA and Canada, Baetiscidae were later found in the fossil record as well, specifically in Cretaceous of Brazil and Eocene Baltic amber. An enigmatic piece of fossil evidence are two larvae from the Early Cretaceous Koonwarra Fossil Bed in Australia, which have been presumed as attributable to Baetiscidae and briefly discussed in previous studies. In the present contribution, we reinvestigate these fossils and confirm their attribution to the family Baetiscidae. These larvae are depicted and described as Koonwarrabaetisca jelli gen. et sp. nov. and Koonwarrabaetisca duncani sp. nov. For both Cretaceous genera Protobaetisca Staniczek, 2007 and Koonwarrabaetisca gen. nov. we establish a new subfamily Protobaetiscinae subfam. nov. within the family Baetiscidae, based on the presence of markedly shortened thoracic sterna. The phylogenetic position of newly described subfamily is clarified using a cladistic analysis; Protobaetiscinae subfam. nov. forms a monophyletic clade, sister to Baetiscinae. The confirmation of the distribution of Baetiscidae in the Cretaceous of Australia suggests almost worldwide distribution of this family in the deep time. Given their limited dispersal abilities, this distributional pattern can be best explained by the Pangean origin for this family, moving the time of their origin at least to the Early Jurassic. The larvae of Koonwarrabaetisca gen. nov. exhibit the same ecomorphological specialization as the rest of Baetiscidae, that supporting with a high probability their lifestyle similar to extant Baetisca Walsh, 1862. The larvae probably lived in the flowing water with stony substrate densely covered by filamentous algae, and in the places of accumulation of dead plant and algae matter during the last instars. Thus, Koonwarrabaetisca gen. nov. could be the allochthonous component in mayfly fauna of the Koonwarra paleolake.

Middle Jurassic insect mines on gymnosperms provide missing links to early mining evolution.

We investigated the mining mode of insect feeding, involving larval consumption of a plant's internal tissues, from the Middle Jurassic (165 million years ago) Daohugou locality of Northeastern China. Documentation of mining from the Jurassic Period is virtually unknown, and results from this time interval would address mining evolution during the temporal gap of mine-seed plant diversifications from the previous Late Triassic to the subsequent Early Cretaceous. Plant fossils were examined with standard microscopic procedures for herbivory and used the standard functional feeding group-damage-type system of categorizing damage. All fossil mines were photographed and databased. We examined 2014 plant specimens, of which 27 occurrences on 14 specimens resulted in eight, new, mine damage types (DTs) present on six genera of bennettitalean, ginkgoalean, and pinalean gymnosperms. Three conclusions emerge from this study. First, these mid-Mesozoic mines are morphologically conservative and track plant host anatomical structure rather than plant phylogeny. Second, likely insect fabricators of these mines were three basal lineages of polyphagan beetles, four basal lineages of monotrysian moths, and a basal lineage tenthredinoid sawflies. Third, the nutrition hypothesis, indicating that miners had greater access to nutritious, inner tissues of new plant lineages, best explains mine evolution during the mid-Mesozoic.

Outside the pattern: Evolution of the genital asymmetry in Saicinae (Hemiptera: Heteroptera: Reduviidae).

Despite genital structures in insects being consistently important as systematic and taxonomy evidence, within assassin bugs (Heteroptera, Reduviidae) at least, the male and female genitalic structures of several subfamilies are poorly or totally unknown. The genital structure is mostly symmetric within Saicinae genera, but male genital asymmetry has been recorded in Gallobelgicus, Polytoxus, and recently Pseudosaica. The subfamily has been considered as closely related to Emesinae and Visayanocorinae but this hypothesis has never been tested using a comprehensive taxa sampling, being a constraint to test morphological traits changes or exploring characters' evolution hypotheses. Here, we compiled a morphological data set of 170 characters that includes external morphological characters and genitalia of both sexes of Saicinae which was analyzed cladistically including 55 terminals, comprising 16 genera (64% of the generic diversity), 43 species of Saicinae and 12 outgroups. Saicinae was recovered as polyphyletic, Saicireta correntina is recovered as sister-species of Empicoris armatus + Collartida (Emesinae), Oncerotrachelus, Carayonia (Visayanocorinae), and the Clade Saicinae sensu stricto. Carayonia orientalis is recovered as sister-species of Saicinae sensu stricto. Ancestral state reconstruction of symmetry of the male genitalia shows an ancestor with symmetric male genitalia, two independent emergences of asymmetrical male genitalia within Saicinae sensu stricto, and the asymmetrical endosomal sclerites appearing before the other asymmetric traits.

Description of the larva of Macrostemum floridum (Navás 1929) (Trichoptera: Hydropsychidae) and its feeding habits in Kaengkrung National Park, southern Thailand.

A morphological description of the last (fifth) instar larva of Macrostemum floridum (Navás 1929), its life cycle, and feeding habits are provided. The study was conducted along the Khlong Yan Stream in Kaengkrung National Park, southern Thailand. Larval identification was achieved by the metamorphotype method of associating immature stages with the adult. A graph of head capsule widths for all instars is provided. Gut content analysis showed that the main food items were diatoms, green algae, and blue-green algae. The larvae construct fixed retreats and feed as collecting filterers.

The neuronal innervation pattern of the subgenual organ complex in Peruphasma schultei (Insecta: Phasmatodea).

The proximal tibia of orthopteroid insects contains sensory organs, the subgenual organ complex, detecting mechanical stimuli including substrate vibration. In stick insects, two chordotonal organs occur in close proximity, the subgenual organ and the distal organ, which likely detect substrate vibrations. In most stick insects, both organs are innervated by separate nerve branches. To obtain more data on the neuroanatomy of the subgenual organ complex from the New World phasmids (Occidophasmata), the present study documents the neuronal innervation of sensory organs in the subgenual organ complex of Peruphasma schultei, the first species from Pseudophasmatinae investigated for this sensory complex. The innervation pattern shows a distinct nerve branch for the subgenual organ and for the distal organ in most cases. Some variability in the innervation, which generally occurs for these chordotonal organs, was noted for both organs in P. schultei. The most common innervation for both organs was by a single nerve branch for each organ. The innervation of the subgenual organ resembled the nerve pattern of another New World phasmid, but was simpler than in the Old World phasmids (Oriophasmata) studied so far. Therefore, the peripheral neuronal innervation of sensory organs could reflect phylogenetic relationships and provide phylogenetic information, while the overall neuroanatomy of the subgenual organ complex is similar in stick insects.