Lower Ordovician synziphosurine reveals early euchelicerate diversity and evolution.

Euchelicerata is a clade of arthropods comprising horseshoe crabs, scorpions, spiders, mites and ticks, as well as the extinct eurypterids (sea scorpions) and chasmataspidids. The understanding of the ground plans and relationships between these crown-group euchelicerates has benefited from the discovery of numerous fossils. However, little is known regarding the origin and early evolution of the euchelicerate body plan because the relationships between their Cambrian sister taxa and synziphosurines, a group of Silurian to Carboniferous stem euchelicerates with chelicerae and an unfused opisthosoma, remain poorly understood owing to the scarce fossil record of appendages. Here we describe a synziphosurine from the Lower Ordovician (ca. 478 Ma) Fezouata Shale of Morocco. This species possesses five biramous appendages with stenopodous exopods bearing setae in the prosoma and a fully expressed first tergite in the opisthosoma illuminating the ancestral anatomy of the group. Phylogenetic analyses recover this fossil as a member of the stem euchelicerate family Offacolidae, which is characterized by biramous prosomal appendages. Moreover, it also shares anatomical features with the Cambrian euarthropod Habelia optata, filling the anatomical gap between euchelicerates and Cambrian stem taxa, while also contributing to our understanding of the evolution of euchelicerate uniramous prosomal appendages and tagmosis.

Discovery of diverse Pectocaris species at the Cambrian series 2 Hongjingshao formation Xiazhuang section (Kunming, SW China) and its ecological, taphonomic, and biostratigraphic implications.

Pectocaris species are intermediate- to large-sized Cambrian bivalved arthropods. Previous studies have documented Pectocaris exclusively from the Cambrian Series 2 Stage 3 Chengjiang biota in Yu'anshan Formation, Chiungchussu Stage in SW China. In this study, we report Pectocaris paraspatiosa sp. nov., and three other previously known Pectocaris from the Xiazhuang section in Kunming, which belongs to the Hongjingshao Formation and is a later phase within Cambrian Stage 3 than the Yu'anshan Formation. The new species can be distinguished from its congeners by the sparsely arranged endopodal endites and the morphologies of the abdomen, telson, and telson processes. We interpret P. paraspatiosa sp. nov. as a filter-feeder and a powerful swimmer adapted to shallow, agitated environments. Comparison among the Pectocaris species reinforces previous views that niche differentiation had been established among the congeneric species based on morphological differentiation. Our study shows the comprehensive occurrences of Pectocaris species outside the Chengjiang biota for the first time. With a review of the shared fossil taxa of Chengjiang and Xiaoshiba biotas, we identify a strong biological connection between the Yu'anshan and Hongjingshao Formations.

Unveiling the ventral morphology of a rare early Cambrian great appendage arthropod from the Chengjiang biota of China.

BACKGROUND: The early Cambrian arthropod clade Megacheira, also referred to as great appendage arthropods, comprised a group of diminutive and elongated predators during the early Palaeozoic era, around 518 million years ago. In addition to those identified in the mid-Cambrian Burgess Shale biota, numerous species are documented in the renowned 518-million-year-old Chengjiang biota of South China. Notably, one species, Tanglangia longicaudata, has remained inadequately understood due to limited available material and technological constraints. In this study, we, for the first time, examined eight fossil specimens (six individuals) utilizing state-of-the-art µCT and computer-based 3D rendering techniques to unveil the hitherto hidden ventral and appendicular morphology of this species. RESULTS: We have identified a set of slender endopodites gradually narrowing distally, along with a leaf-shaped exopodite adorned with fringed setae along its margins, and a small putative exite attached to the basipodite. Our techniques have further revealed the presence of four pairs of biramous appendages in the head, aligning with the recently reported six-segmented head in other early euarthropods. Additionally, we have discerned two peduncle elements for the great appendage. These findings underscore that, despite the morphological diversity observed in early euarthropods, there exists similarity in appendicular morphology across various groups. In addition, we critically examine the existing literature on this taxon, disentangling previous mislabelings, mentions, descriptions, and, most importantly, illustrations. CONCLUSIONS: The µCT-based investigation of fossil material of Tanglangia longicaudata, a distinctive early Cambrian euarthropod from the renowned Chengjiang biota, enhances our comprehensive understanding of the evolutionary morphology of the Megacheira. Its overall morphological features, including large cup-shaped eyes, raptorial great appendages, and a remarkably elongated telson, suggest its potential ecological role as a crepuscular predator and adept swimmer in turbid waters.

New Canadian amber deposit fills gap in fossil record near end-Cretaceous mass extinction.

Amber preserves an exceptional record of tiny, soft-bodied organisms and chemical environmental signatures, elucidating the evolution of arthropod lineages and the diversity, ecology, and biogeochemistry of ancient ecosystems. However, globally, fossiliferous amber deposits are rare in the latest Cretaceous and surrounding the Cretaceous-Paleogene (K-Pg) mass extinction.1,2,3,4,5 This faunal gap limits our understanding of arthropod diversity and survival across the extinction boundary.2,6 Contrasting hypotheses propose that arthropods were either relatively unaffected by the K-Pg extinction or experienced a steady decline in diversity before the extinction event followed by rapid diversification in the Cenozoic.2,6 These hypotheses are primarily based on arthropod feeding traces on fossil leaves and time-calibrated molecular phylogenies, not direct observation of the fossil record.2,7 Here, we report a diverse amber assemblage from the Late Cretaceous (67.04 ± 0.16 Ma) of the Big Muddy Badlands, Canada. The new deposit fills a critical 16-million-year gap in the arthropod fossil record spanning the K-Pg mass extinction. Seven arthropod orders and at least 11 insect families have been recovered, making the Big Muddy amber deposit the most diverse arthropod assemblage near the K-Pg extinction. Amber chemistry and stable isotopes suggest the amber was produced by coniferous (Cupressaceae) trees in a subtropical swamp near remnants of the Western Interior Seaway. The unexpected abundance of ants from extant families and the virtual absence of arthropods from common, exclusively Cretaceous families suggests that Big Muddy amber may represent a yet unsampled Late Cretaceous environment and provides evidence of a faunal transition before the end of the Cretaceous.

A morphofunctional study of the jumping apparatus in globular springtails.

Springtails are notable for their jumping apparatus and latch-mediated spring mechanism. The challenge, in the light of the tiny size and rapid movement of these organisms, has been to understand the morphological intricacies of this spring system. This study takes an approach that integrates SEM, MicroCT, cLSM and high-speed video recordings to understand the composition and functionality of the jumping apparatus in Megalothorax minimus (Neelipleona), Dicyrtomina ornata and Dicyrtomina minuta (Symphypleona). We focus on reconstructing, describing, and understanding the functioning of structures such as basal plates, musculature and furca. The dimensions of the jumping apparatus in Dicyrtomina and Megalothorax differ significantly from those in elongated springtails. A hypothesis of functional coherence between taxa, based on muscle connections and basal plates, is postulated. High-speed video recordings provide information on: 1) furca release timing and function during jumping and self-righting; 2) performance properties of manubrium, dens and mucro in interaction with the ground and in take-off; 3) possible pre-release furca moves. The study underscores the need for further research employing a variety of visualization methods in order to explore additional aspects such as retinaculum unlatching and furca flexion/extension muscles.

A network analysis of early arthropod evolution and the potential of the primitive.

It is often thought that the primitive is simpler, and that the complex is generated from the simple by some process of self-assembly or self-organization, which ultimately consists of the spontaneous and fortuitous collision of elementary units. This idea is included in the Darwinian theory of evolution, to which is added the competitive mechanism of natural selection. To test this view, we studied the early evolution of arthropods. Twelve groups of arthropods belonging to the Burgess Shale, Orsten Lagerstätte, and extant primitive groups were selected, their external morphology abstracted and codified in the language of network theory. The analysis of these networks through different network measures (network parameters, topological descriptors, complexity measures) was used to carry out a Principal Component Analysis (PCA) and a Hierarchical Cluster Analysis (HCA), which allowed us to obtain an evolutionary tree with distinctive/novel features. The analysis of centrality measures revealed that these measures decreased throughout the evolutionary process, and led to the creation of the concept of evolutionary developmental potential. This potential, which measures the capacity of a morphological unit to generate changes in its surroundings, is concomitantly reduced throughout the evolutionary process, and demonstrates that the primitive is not simple but has a potential that unfolds during this process. This means for us the first empirical evolutionary evidence of our theory of evolution as a process of unfolding.

Rapid divergent evolution of internal female genitalia and the coevolution of male genital morphology revealed by micro-computed tomography.

Animal genitalia are thought to evolve rapidly and divergently in response to sexual selection. Studies of genital evolution have focused largely on male genitalia. The paucity of work on female genital morphology is probably due to problems faced in quantifying shape variation, due to their composition and accessibility. Here we use a combination of micro-computed tomography, landmark free shape quantification and phylogenetic analysis to quantify the rate of female genital shape evolution among 29 species of Antichiropus millipedes, and their coevolution with male genitalia. We found significant variation in female and male genital shape among species. Male genital shape showed a stronger phylogenetic signal than female genital shape, although the phylogenetic signal effect sizes did not differ significantly. Male genital shape was found to be evolving 1.2 times faster than female genital shape. Female and male genital shape exhibited strong correlated evolution, indicating that genital shape changes in one sex are associated with corresponding changes in the genital shape of the other sex. This study adds novel insight into our growing understanding of how female genitalia can evolve rapidly and divergently, and highlights the advantages of three-dimensional techniques and multivariate analyses in studies of female genital evolution.

Convergent evolution of ventral adaptations for enrolment in trilobites and extant euarthropods.

The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott-Rust Quarry in New York State, USA. Walcott-Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages.

Uniquely preserved gut contents illuminate trilobite palaeophysiology.

Trilobites are among the most iconic of fossils and formed a prominent component of marine ecosystems during most of their 270-million-year-long history from the early Cambrian period to the end Permian period1. More than 20,000 species have been described to date, with presumed lifestyles ranging from infaunal burrowing to a planktonic life in the water column2. Inferred trophic roles range from detritivores to predators, but all are based on indirect evidence such as body and gut morphology, modes of preservation and attributed feeding traces; no trilobite specimen with internal gut contents has been described3,4. Here we present the complete and fully itemized gut contents of an Ordovician trilobite, Bohemolichas incola, preserved three-dimensionally in a siliceous nodule and visualized by synchrotron microtomography. The tightly packed, almost continuous gut fill comprises partly fragmented calcareous shells indicating high feeding intensity. The lack of dissolution of the shells implies a neutral or alkaline environment along the entire length of the intestine supporting digestive enzymes comparable to those in modern crustaceans or chelicerates. Scavengers burrowing into the trilobite carcase targeted soft tissues below the glabella but avoided the gut, suggesting noxious conditions and possibly ongoing enzymatic activity.

The early Cambrian Kylinxia zhangi and evolution of the arthropod head.

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 described1), 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 euarthropods6,7,8,9,10 is bolstered by morphological similarities and inferred phylogenetic continuity between Kylinxia and other stem-group euarthropods.

Raptorial appendages of the Cambrian apex predator Anomalocaris canadensis are built for soft prey and speed.

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.

A new species of Ptenothrix Börner, 1906 (Collembola, Symphypleona, Dicyrtomidae) from South Korea.

A new species of Collembola, Ptenothrix koreanensis sp. nov., is described and illustrated based on specimens from South Korea. It was found in a coniferous forest dominated by the Korean pine Pinus koraiensis Siebold & Zuccarini. Ptenothrix koreanensis sp. nov. is characterized by a peculiar color pattern in which most of its median dorsal side of the large abdomen is plain milky white. This new species also has the formula of 1, 1, 2, 2, 1, 3 of median spiny chaetae on the clypeal area of the head, and the formula of 6, 5, 5, 4 of prelabral and labral chaetae. These characteristics make it easy to distinguish the new species from other two possibly related species, P. ciliophora Yosii & Lee and P. saxatilis Yosii & Lee. Apart from these characteristics, the new species can also be separated from another possibly related species, P. higashihirajii Yosii, by the morphology of ps1 and pi2 chaetae on the small abdomen of the female. An identification key for the East Asian species of Ptenothrix is also provided.

Comment on "The lower Cambrian lobopodian Cardiodictyon resolves the origin of euarthropod brains".

Strausfeld et al. (Report, 24 Nov 2022, p. 905) claim that Cambrian fossilized nervous tissue supports the interpretation that the ancestral panarthropod brain was tripartite and unsegmented. We argue that this conclusion is unsupported, and developmental data from living onychophorans contradict it.

Response to Comment on "The lower Cambrian lobopodian Cardiodictyon resolves the origin of euarthropod brains".

Budd et al. challenge the identity of neural traces reported for the Cambrian lobopodian Cardiodictyon catenulum. Their argumentation is unsupported, as are objections with reference to living Onychophora that misinterpret established genomic, genetic, developmental, and neuroanatomical evidence. Instead, phylogenetic data corroborate the finding that the ancestral panarthropod head and brain is unsegmented, as in C. catenulum.

Life cycle evolution in the trilobites Balangia and Duyunaspis from the Cambrian Series 2 (Stage 4) of South China.

The evolution process can be reconstructed by tracking the changes in the dynamic characters of life cycles. A number of related trilobites from the Cambrian of South China provide additional information for the study of trilobite evolutionary patterns, which has been hampered by previous incomplete fossil record though. Here, Balangia and Duyunaspis represent related Cambrian oryctocephalid trilobites from South China, are comprehensively discussed over the ontogeny, and the results show that, from B. balangensis via D. duyunensis to D. jianheensis, their exoskeletal morphology shows a directional evolution. Based on the direction of evolutionary changes in the development of Balangia and Duyunaspis, we speculate that Duyunaspis likely evolved from Balangia instead of Balangia evolved from Duyunaspis, as was previously assumed. This inference is also supported by the phylogenetic tree. This research provides not only a better understanding of the mechanisms of evolution in trilobites, but also new insights for the relationship between developmental evolutionary changes and phylogeny in trilobites.

Three-dimensional morphology of the biramous appendages in Isoxys from the early Cambrian of South China, and its implications for early euarthropod evolution.

Early euarthropod evolution involved a major transition from lobopodian-like taxa to organisms featuring a segmented, well-sclerotized trunk (arthrodization) and limbs (arthropodization). However, the precise origin of a completely arthrodized trunk and arthropodized ventral biramous appendages remain controversial, as well as the early onset of anterior-posterior limb differentiation in stem-group euarthropods. New fossil material and micro-computed tomography inform the detailed morphology of the arthropodized biramous appendages in the carapace-bearing euarthropod Isoxys curvirostratus from the early Cambrian Chengjiang biota. In addition to well-developed grasping frontal appendages, I. curvirostratus possesses two batches of morphologically and functionally distinct biramous limbs. The first batch consists of four pairs of short cephalic appendages with robust endites with a feeding function, whereas the second batch has more elongate trunk appendages for locomotion. Critically, our new material shows that the trunk of I. curvirostratus was not arthrodized. The results of our phylogenetic analyses recover isoxyids as some of the earliest branching sclerotized euarthropods, and strengthens the hypothesis that arthropodized biramous appendages evolved before full body arthrodization.

Shared patterns of segment size development in trilobites and vertebrates.

The relative sizes of body segments are a major determinant of the shape and functionality of an animal. Developmental biases affecting this trait can therefore have major evolutionary implications. In vertebrates, a molecular activator/inhibitor mechanism, known as the inhibitory cascade (IC), produces a simple and predictable pattern of linear relative size along successive segments. The IC model is considered the default mode of vertebrate segment development and has produced long-term biases in the evolution of serially homologous structures such as teeth, vertebrae, limbs, and digits. Here we investigate whether the IC model or an IC-like model also has controls on segment size development in an ancient and hyperdiverse group of extinct arthropods, the trilobites. We examined segment size patterning in 128 trilobite species, and during ontogenetic growth in three trilobite species. Linear relative segment size patterning is prominent throughout the trunk of trilobites in the adult form, and there is strict regulation of this patterning in newly developing segments in the pygidium. Extending the analysis to select stem and modern arthropods suggests that the IC is a common default mode of segment development capable of producing long-term biases in morphological evolution across arthropods as it does in vertebrates.

The unbearable uncertainty of panarthropod relationships.

Panarthropoda, the clade comprising the phyla Onychophora, Tardigrada and Euarthropoda, encompasses the largest majority of animal biodiversity. The relationships among the phyla are contested and resolution is key to understanding the evolutionary assembly of panarthropod bodyplans. Molecular phylogenetic analyses generally support monophyly of Onychophora and Euarthropoda to the exclusion of Tardigrada (Lobopodia hypothesis), which is also supported by some analyses of morphological data. However, analyses of morphological data have also been interpreted to support monophyly of Tardigrada and Euarthropoda to the exclusion of Onychophora (Tactopoda hypothesis). Support has also been found for a clade of Onychophora and Tardigrada that excludes Euarthropoda (Protarthropoda hypothesis). Here we show, using a diversity of phylogenetic inference methods, that morphological datasets cannot discriminate statistically between the Lobopodia, Tactopoda and Protarthropoda hypotheses. Since the relationships among the living clades of panarthropod phyla cannot be discriminated based on morphological data, we call into question the accuracy of morphology-based phylogenies of Panarthropoda that include fossil species and the evolutionary hypotheses based upon them.

Interpreting fossilized nervous tissues.

Paleoneuranatomy is an emerging subfield of paleontological research with great potential for the study of evolution. However, the interpretation of fossilized nervous tissues is a difficult task and presently lacks a rigorous methodology. We critically review here cases of neural tissue preservation reported in Cambrian arthropods, following a set of fundamental paleontological criteria for their recognition. These criteria are based on a variety of taphonomic parameters and account for morphoanatomical complexity. Application of these criteria shows that firm evidence for fossilized nervous tissues is less abundant and detailed than previously reported, and we synthesize here evidence that has stronger support. We argue that the vascular system, and in particular its lacunae, may be central to the understanding of many of the fossilized peri-intestinal features known across Cambrian arthropods. In conclusion, our results suggest the need for caution in the interpretation of evidence for fossilized neural tissue, which will increase the accuracy of evolutionary scenarios. Also see the video abstract here: https://youtu.be/2_JlQepRTb0.

Sexually dimorphic characters of the ultimate legs in lithobiid centipedes (Myriapoda, Chilopoda, and Lithobiomorpha): Morphology and implications for reproductive behavior.

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.