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The cuticle is a key evolutionary innovation that played a crucial role in arthropod terrestrialization. Extensive research has elucidated the chemical and structural composition of the cuticle in extant arthropods, while fossil studies have further informed our understanding of cuticle evolution. This study examines the three-dimensionally preserved cuticular structure of the Early Devonian trigonotarbid arachnid genus Palaeocharinus, from the Rhynie chert of Scotland (∼408 Ma). Trigonotarbids, an extinct group of tetrapulmonate arachnids, are among the earliest known unequivocally terrestrial arthropods, and thus may shed light on the evolution of terrestriality. Using high-resolution Confocal Laser Scanning Microscopy (CLSM), we reveal detailed morphological features at the nanometre level. The external cuticle surface of Palaeocharinus is characterized by polygonal scales, sensilla, and small pores identified as the openings of dermal glands and wax canals. Internally, the cuticle exhibits polygonal clusters of pore canals, through which wax was transported from the epidermis to the cuticular surface. The pore canals twist along their vertical axes, reflecting the "twisted plywood" or Bouligand arrangement of chitin-protein microfibril planes characteristic of modern arthropod cuticles. Overall, the cuticle of Palaeocharinus is characteristically thick relative to those of other extinct and extant chelicerates, such thickening being a possible adaptation to terrestrial life., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Background: The sea spiders (Pycnogonida Latreille, 1810) of the Hunsrück Slate (Lower Devonian, ~400 million years ago) are iconic in their abundance, exquisite pyritic preservation, and in their distinctive body plan compared to extant sea spiders (Pantopoda Gerstäcker, 1863). Consequently, the Hunsrück sea spiders are important in understanding the deep evolutionary history of Pycnogonida, yet they remain poorly characterised, impacting upon attempts to establish a time-calibrated phylogeny of sea spiders., Methods: Here, we investigated previously described and new material representing four of the five Hunsrück pycnogonids: Flagellopantopus blocki Poschmann & Dunlop, 2006; Palaeoisopus problematicus Broili, 1928; Palaeopantopus maucheri Broili, 1929; and Pentapantopus vogteli Kühl, Poschmann & Rust, 2013; as well as a few unidentified specimens. Using X-ray microtomography and Reflectance Transformation Imaging, we describe new fossils, provide evidence for newly revealed anatomical features, and interpret these data in comparison to extant species. We also reinterpret the previously published illustration of the (probably lost) holotype of Palaeothea devonica Bergström, Stürmer & Winter, 1980., Results: We provide the first detailed description of the cephalic appendages of Palaeoisopus problematicus and revise the interpretation of the organisation of its ocular tubercle. Furthermore, we provide new insights into the structure of the legs and the proboscis of Palaeopantopus maucheri , the first description of the body of Flagellopantopus blocki and describe a new specimen of Pentapantopus vogteli , demonstrating that it had eight legs, in contrast to previous interpretations. We argue that, contrary to previous suggestions, Palaeothea devonica probably had a different body plan from extant pantopods. We discuss the ecological traits of the Hunsrück pycnogonids based on their morphological adaptations, and conclude that there is no compelling evidence of Pantopoda in the Devonian. Through comparative interpretation of the legs as well as general morphology, we can divide the Hunsrück pycnogonids into two morphological groups, while Pantopoda constitutes a third morphological group., Competing Interests: The authors declare that they have no competing interests., (© 2024 Sabroux et al.)

The Carboniferous myriapod Arthropleura is the largest arthropod of all time, but its fossils are usually incomplete, limiting the understanding of its anatomy, ecology, and relationships. Micro-computed tomography applied to exceptionally preserved specimens from the Carboniferous Montceau-les-Mines Lagerstätte (France) reveals unprecedented details of its functional anatomy, such as the head and mouthparts. Arthropleura shares features with both millipedes and centipedes. Total-evidence phylogeny combining morphological and transcriptomic data resolves Arthropleura alone as a stem group millipede, but the inclusion of the highly incomplete Siluro-Devonian Eoarthropleura draws it deeper into the myriapod stem. Arthropleura suggests transitional morphology between clades united primarily by molecular information and underscores the value of total-evidence phylogenetics to understanding evolutionary history.

Recent advances in higher-level invertebrate phylogeny have leveraged shared features of genomic architecture to resolve contentious nodes across the tree of life. Yet, the interordinal relationships within Chelicerata have remained recalcitrant given competing topologies in recent molecular analyses. As such, relationships between topologically unstable orders remain supported primarily by morphological cladistic analyses. Solifugae, one such unstable chelicerate order, has long been thought to be the sister group of Pseudoscorpiones, forming the clade Haplocnemata, on the basis of eight putative morphological synapomorphies. The discovery, however, of a shared whole genome duplication placing Pseudoscorpiones in Arachnopulmonata provides the opportunity for a simple litmus test evaluating the validity of Haplocnemata. Here, we present the first developmental transcriptome of a solifuge (Titanopuga salinarum) and survey copy numbers of the homeobox genes for evidence of systemic duplication. We find that over 70% of the identified homeobox genes in T. salinarum are retained in a single copy, while representatives of the arachnopulmonates retain orthologs of those genes as two or more copies. Our results refute the placement of Solifugae in Haplocnemata. Subsequent reevaluation of putative interordinal morphological synapomorphies among chelicerates reveals a high incidence of homoplasy, reversals, and inaccurate coding within Haplocnemata and other small clades, as well as Arachnida more broadly, suggesting existing morphological character matrices are insufficient to resolve chelicerate phylogeny., (© 2023 The Authors. Evolution & Development published by Wiley Periodicals LLC.)

Knowledge of Cambrian animal anatomy is limited by preservational processes that result in compaction, size bias, and incompleteness. We documented pristine three-dimensional (3D) anatomy of trilobites fossilized through rapid ash burial from a pyroclastic flow entering a shallow marine environment. Cambrian ellipsocephaloid trilobites from Morocco are articulated and undistorted, revealing exquisite details of the appendages and digestive system. Previously unknown anatomy includes a soft-tissue labrum attached to the hypostome, a slit-like mouth, and distinctive cephalic feeding appendages. Our findings resolve controversy over whether the trilobite hypostome is the labrum or incorporates it and establish crown-group euarthropod homologies in trilobites. This occurrence of moldic fossils with 3D soft parts highlights volcanic ash deposits in marine settings as an underexplored source for exceptionally preserved organisms.

The early Cambrian Kylinxia zhangi occupies a pivotal position in arthropod evolution, branching from the euarthropod stem lineage between radiodonts (Anomalocaris and relatives) and "great-appendage" arthropods. 1 , 2 Its combination of appendage and exoskeletal features is viewed as uniquely bridging the morphologies of so-called "lower" and "upper" stem-group euarthropods. 3 , 4 Microtomographic study of new specimens of Kylinxia refines and corrects previous interpretation of head structures in this species. Phylogenetic analyses incorporating new data reinforce the placement of Kylinxia in the euarthropod stem group but support new hypotheses of head evolution. The head of Kylinxia is composed of six segments, as in extant mandibulates, e.g., insects. 5 In Kylinxia, these are an anterior sclerite associated with an unpaired median eye and paired lateral eyes (thus three rather than five eyes as was previously described 1 ), deutocerebral frontal-most appendages, and four pairs of biramous appendages (rather than two pairs of uniramous appendages). Phylogenetic trees suggest that a six-segmented head in the euarthropod crown group was already acquired by a common ancestor with Kylinxia. The segmental alignment and homology of spinose frontal-most appendages between radiodonts and upper stem-group euarthropods 6 , 7 , 8 , 9 , 10 is bolstered by morphological similarities and inferred phylogenetic continuity between Kylinxia and other stem-group euarthropods., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

We present a genome assembly from an individual male Strigamia acuminata (centipede; Arthropoda; Chilopoda; Geophilomorpha; Geophilidae; Linotaeniinae). The genome sequence is 237.5 megabases in span. Most of the assembly is scaffolded into 11 chromosomal pseudomolecules, including the X and Y sex chromosomes. The mitochondrial genome has also been assembled and is 15.07 kilobases in length., Competing Interests: No competing interests were disclosed., (Copyright: © 2023 Edgecombe GD et al.)

Aim: To investigate the drivers of intra-specific genetic diversity in centipedes, a group of ancient predatory soil arthropods., Location: Asia, Australasia and Europe., Time Period: Present., Major Taxa Studied: Centipedes (Class: Chilopoda)., Methods: We assembled a database of 1245 mitochondrial cytochrome c oxidase subunit I sequences representing 128 centipede species from all five orders of Chilopoda. This sequence dataset was used to estimate genetic diversity for centipede species and compare its distribution with estimates from other arthropod groups. We studied the variation in centipede genetic diversity with species traits and biogeography using a beta regression framework, controlling for the effect of shared evolutionary history within a family., Results: A wide variation in genetic diversity across centipede species (0-0.1713) falls towards the higher end of values among arthropods. Overall, 27.57% of the variation in mitochondrial COI genetic diversity in centipedes was explained by a combination of predictors related to life history and biogeography. Genetic diversity decreased with body size and latitudinal position of sampled localities, was greater in species showing maternal care and increased with geographic distance among conspecifics., Main Conclusions: Centipedes fall towards the higher end of genetic diversity among arthropods, which may be related to their long evolutionary history and low dispersal ability. In centipedes, the negative association of body size with genetic diversity may be mediated by its influence on local abundance or the influence of ecological strategy on long-term population history. Species with maternal care had higher genetic diversity, which goes against expectations and needs further scrutiny. Hemispheric differences in genetic diversity can be due to historic climatic stability and lower seasonality in the southern hemisphere. Overall, we find that despite the differences in mean genetic diversity among animals, similar processes related to life-history strategy and biogeography are associated with the variation within them., Competing Interests: Conflict of Interest Statement The authors declare no conflict of interest.

The stem-group euarthropod Anomalocaris canadensis is one of the largest Cambrian animals and is often considered the quintessential apex predator of its time. This radiodont is commonly interpreted as a demersal hunter, responsible for inflicting injuries seen in benthic trilobites. However, controversy surrounds the ability of A. canadensis to use its spinose frontal appendages to masticate or even manipulate biomineralized prey. Here, we apply a new integrative computational approach, combining three-dimensional digital modelling, kinematics, finite-element analysis (FEA) and computational fluid dynamics (CFD) to rigorously analyse an A. canadensis feeding appendage and test its morphofunctional limits. These models corroborate a raptorial function, but expose inconsistencies with a capacity for durophagy. In particular, FEA results show that certain parts of the appendage would have experienced high degrees of plastic deformation, especially at the endites, the points of impact with prey. The CFD results demonstrate that outstretched appendages produced low drag and hence represented the optimal orientation for speed, permitting acceleration bursts to capture prey. These data, when combined with evidence regarding the functional morphology of its oral cone, eyes, body flaps and tail fan, suggest that A. canadensis was an agile nektonic predator that fed on soft-bodied animals swimming in a well-lit water column above the benthos. The lifestyle of A. canadensis and that of other radiodonts, including plausible durophages, suggests that niche partitioning across this clade influenced the dynamics of Cambrian food webs, impacting on a diverse array of organisms at different sizes, tiers and trophic levels.

Many species of lithobiomorph centipedes present a pronounced sexual dimorphism reflected in remarkable structural modifications on the ultimate legs of males. Most records of these male secondary sexual characters addressed taxonomy, helping to identify and characterize species or diagnose genera, but information on their diversity, detailed morphology and possible function(s) is scarce. In this study, nine species of the two lithobiid genera Lithobius Leach, 1814 and Eupolybothrus Verhoeff, 1907 were investigated, using light and scanning electron microscopy to document the detailed morphology of secondary sexual characters of male ultimate legs. Secondary sexual characters affecting the cuticle of the ultimate legs are described in detail and found to often be associated with sensilla, interpreted here as chemo- and mechanoreceptors, and with clusters of pores, a hitherto undescribed pore-distribution for this group. The tibial nodule of the species Lithobius nodulipes Latzel, 1880, was additionally examined with histological semi-thin sections. These results revealed that the clustered pores are connected to glandular tissue, and are, based on their morphology, interpreted as openings of flexo-canal epidermal glands. The presence of various sensory and glandular structures associated with sexual dimorphism indicates a likely role during courtship and mating. The closely related species examined in this research show comparable dimorphic structures, which are otherwise species-specific. Morphological observations on secondary sexual structures inform on reproductive biology in groups like lithobiomorphs for which there are limited behavioral data., (© 2022 Wiley Periodicals LLC.)

Recent transcriptomic studies of myriapod phylogeny have been based on relatively small datasets with <40 myriapod terminals and variably supported or contradicted the traditional morphological groupings of Progoneata and Dignatha. Here we amassed a large dataset of 104 myriapod terminals, including multiple species for each of the four myriapod classes. Across the tree, most nodes are stable and well supported. Most analyses across a range of gene occupancy levels provide moderate to strong support for a deep split of Myriapoda into Symphyla + Pauropoda (=Edafopoda) and an uncontradicted grouping of Chilopoda + Diplopoda (=Pectinopoda nov.), as in other recent transcriptome-based analyses; no analysis recovers Progoneata or Dignatha as clades. As in all recent multi-locus and phylogenomic studies, chilopod interrelationships resolve with Craterostigmus excluded from Amalpighiata rather than uniting with other centipedes with maternal brood care in Phylactometria. Diplopod ordinal interrelationships are largely congruent with morphology-based classifications. Chilognathan clades that are not invariably advocated by morphologists include Glomerida + Glomeridesmida, such that the volvation-related characters of pill millipedes may be convergent, and Stemmiulida + Polydesmida more closely allied to Juliformia than to Callipodida + Chordeumatida. The latter relationship implies homoplasy in spinnerets and contradicts Nematophora. A time-tree with nodes calibrated by 25 myriapod and six outgroup fossil terminals recovers Cambrian-Ordovician divergences for the deepest splits in Myriapoda, Edafopoda and Pectinopoda, predating the terrestrial fossil record of myriapods as in other published chronograms, whereas age estimates within Chilopoda and Diplopoda overlap with or do not appreciably predate the calibration fossils. The grouping of Chilopoda and Diplopoda is recovered in all our analyses and is formalized as Pectinopoda nov., named for the shared presence of mandibular comb lamellae. New taxonomic proposals for Chilopoda based on uncontradicted clades are Tykhepoda nov. for the three blind families of Scolopendromorpha that share a "sieve-type" gizzard, and Taktikospina nov. for Scolopendromorpha to the exclusion of Mimopidae., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Eurypterids (sea scorpions) are extinct aquatic chelicerates. Within this group, members of Pterygotidae represent some of the largest known marine arthropods. Representatives of this family all have hypertrophied, anteriorly-directed chelicerae and are commonly considered Silurian and Devonian apex predators. Despite a long history of research interest in these appendages, pterygotids have been subject to limited biomechanical investigation. Here, we present finite element analysis (FEA) models of four different pterygotid chelicerae-those of Acutiramus bohemicus , Erettopterus bilobus , Jaekelopterus rhenaniae , and Pterygotus anglicus -informed through muscle data and finite element models (FEMs) of chelae from 16 extant scorpion taxa. We find that Er. bilobus and Pt. anglicus have comparable stress patterns to modern scorpions, suggesting a generalised diet that probably included other eurypterids and, in the Devonian species, armoured fishes, as indicated by co-occurring fauna. Acutiramus bohemicus is markedly different, with the stress being concentrated in the proximal free ramus and the serrated denticles. This indicates a morphology better suited for targeting softer prey. Jaekelopterus rhenaniae exhibits much lower stress across the entire model. This, combined with an extremely large body size, suggests that the species likely fed on larger and harder prey, including heavily armoured fishes. The range of cheliceral morphologies and stress patterns within Pterygotidae demonstrate that members of this family had variable diets, with only the most derived species likely to feed on armoured prey, such as placoderms. Indeed, increased sizes of these forms throughout the mid-Palaeozoic may represent an 'arms race' between eurypterids and armoured fishes, with Devonian pterygotids adapting to the rapid diversification of placoderms., Competing Interests: The authors declare that they have no competing interests., (© 2022 Bicknell et al.)

Among extant animals, Lophotrochozoa accounts for the majority of phyla. 1 This bilaterian clade radiated rapidly during the Cambrian explosion, obfuscating its phylogenetic relationships and rendering many aspects of its early evolution uncertain. Many early lophotrochozoans are known only from isolated skeletal microfossils, "small shelly fossils," often derived from larger animals with complex multi-element skeletons. 2 The discovery of articulated fossils has revealed surprising insights into the animals from which these skeletal pieces were derived, such as paired shells in the mollusc Halkieria. 3 Tommotiids are a key group of phosphatic early skeletal fossils that first appear in the late early Cambrian. 4 , 5 Although their affinities were previously obscure, discoveries of partial scleritomes and investigations of growth and microstructure 6 provide links with Brachiopoda 7 , 8 and Phoronida, 9 two of the lophophorate phyla. By contrast, the body plan of camenellan tommotiids remains a palaeontological mystery, with hypothetical reconstructions representing motile, benthic, dorsally armored worms. 4 , 10 Here, we describe an articulated camenellan (Wufengella bengtsoni gen. et sp. nov.) from the Cambrian Chengjiang Biota, China, revealing the morphology of the scleritome and the first soft tissues from an adult tommotiid. Wufengella carries two dorsal rows of sclerites in a highly asymmetric arrangement, flanked by smaller, cap-shaped sclerites. The scleritome was fringed by iterated fascicles of chaetae and two layers of flattened lobes. Phylogenetic analysis confirms that camenellans occupy a deep branch in lophophorate phylogeny, prior to the acquisition of a sessile lifestyle. Wufengella reveals direct evidence for a metameric body plan reminiscent of annelids early in the evolutionary history of lophophorates. 11 , 12 ., Competing Interests: Declaration of interests O.B. declares possession of shares in FlyDome, the equipment used to generate the RTI datasets., (Copyright © 2022 The Authors. Published by Elsevier Inc. All rights reserved.)

The anterior-most unit of the crown-group arthropod body plan includes three segments, the pre-gnathal segments, that contain three neuromeres that together comprise the brain. Recent work on the development of this anterior region has shown that its three units exhibit many developmental differences to the more posterior segments, to the extent that they should not be considered serial homologs. Building on this revised understanding of the development of the pre-gnathal segments, we suggest a novel scenario for arthropod head evolution. We posit an expansion of an ancestral single-segmented head at the transition from Radiodonta to Deuteropoda in the arthropod stem group. The expanded head subdivided into three segmental units, each maintaining some of the structures of the ancestral head. This scenario is consistent with what we know of head evolution from the fossil record and helps reconcile some of the debates about early arthropod evolution., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)