Enigmatic host-mite relationships: Unraveling the distribution of quill mites on Birds-of-Paradise.

Mites of the family Syringophilidae (Acariformes: Prostigmata: Cheyletoidea) are permanent and obligatory parasites of birds. This study presents an analysis of mite material collected from 22 avian species belonging to the family Paradisaeidae (Passeriformes), revealing the presence of four mite species belonging to four genera: Syringophiloidus attenboroughi n. sp., Peristerophila regiusi n. comb., Picobia frankei, and Gunabopicobia garylarsoni. In the present work, the genus Neoperisterophila is synonymized with the genus Peristerophila. While the genera Syringophiloidus and Picobia were expectedly found on paradisaeid birds, given their prevalence in passerines, the presence of Peristerophila and Gunabopicobia was intriguing, suggesting potential host-switching events. The specificity of these mites varies, with some showing occurrence on hosts of closely related genera and others infesting phylogenetically distant hosts. Notably, the distribution of specific mite species on the Birds-of-Paradise appears to be influenced by both long coevolutionary histories and incidental contacts between often unrelated or intergeneric hybrid species of paradisaeid birds. Furthermore, our research of 104 specimens from 22 Birds-of-Paradise species shows generally low infestation rates across the studied species, suggesting a nuanced interaction between these mites and their avian hosts. Additionally, our network analysis provides a deeper understanding of these host-parasite interactions, revealing a high level of specialization and complexity in these ecological relationships.

Just a matter of size? Evaluating allometry and intersexual heterometry in Pagurus bernhardus using ratios and indices (Decapoda, Anomura).

Heterochely denotes the presence of dissimilarly sized chelipeds on opposite sides of the body, a prevalent occurrence in diverse crustaceans. Conversely, heterometry pertains to the quantifiable disparities in size between these chelipeds. Both chelipeds hold pivotal roles in activities such as foraging, mating, and defense. Consequently, individuals of both genders in heterochelic species exhibit this morphological pattern. Previous studies have identified sexual dimorphism in cheliped size, with males displaying larger major chelipeds compared to females, albeit solely relying on propodus length as a size proxy and focusing solely on the major cheliped. In our study, we meticulously examined 190 specimens of the common European hermit crab Pagurus bernhardus from two collections. We sought to elucidate allometric relationships and assess whether heterometry exhibited sex-based differences when adjusting for body size by using ratios. Our findings revealed that male chelipeds displayed hyperallometric growth relative to females, and all three calculated heterometry indices exhibited significant disparities between the sexes. Consequently, male specimens exhibited larger major and minor chelipeds, even when theoretically matched for body size with females. This phenomenon may be attributed, among other factors, to male-male contests. Should indirect mate selection favor males with larger chelipeds in proportion to their body size, this dynamic could potentiate sexual selection in their favor.

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.

First Records of Picobiine Mites Associated with Birds-of-Paradise: Can Interspecific Sexual Behaviour of Hosts Play a Role in the Distribution of Quill Mite Parasites?

While birds-of-paradise (Passeriformes: Paradisaeidae) are a well-known group of birds, our understanding of their parasites is still limited. This study reports on parasitic quill mites of the subfamily Picobiinae (Acariformes: Syringophilidae), which have never before been recorded on this group of birds. The mite specimens presented in this paper were collected from birds-of-paradise that had been captured in Papua New Guinea and Indonesia in the years 1910-1911 and are now deposited in the Bavarian State Collection of Zoology, Munich, Germany. Two syringophilid species are described as new to science: (i) Picobia frankei sp. n. from the magnificent riflebird Lophorina magnifica, the glossy-mantled manucode Manucodia ater, and the crinkle-collared manucode Manucodia chalybatus, and (ii) Gunabopicobia garylarsoni sp. n. from the twelve-wired bird-of-paradise Seleucidis melanoleucus and the lesser bird-of-paradise Paradisaea minor. We hypothesise that the presence of both picobiine species on phylogenetically unrelated paradisaeids may be caused by the sexual behaviour of these birds, where interspecific copulations may play a role in the switching of parasites between non-closely related host species.

Ventral Morphology of the Non-Trilobite Artiopod Retifacies abnormalis Hou, Chen & Lu, 1989, from the Early Cambrian Chengjiang Biota, China.

The artiopodans represent a diverse group of euarthropods with a typically flattened dorsal exoskeleton that covers numerous pairs of biramous ventral appendages, and which are ubiquitous faunal components of the 518-million-year-old Chengjiang Lagerstätte in South China. Despite their abundance, several Chengjiang artiopodans remain poorly known, such as the large euarthropoda Retifacies abnormalis, Hou, Chen & Lu, 1989, which is distinguished by the presence of mesh-like ornamentation on its dorsal exoskeleton. Although only a few ventral details were described in a single study in 25 years, it has been frequently featured in phylogenetic analyses that explore the relationships between Cambrian euarthropods. Here, we employ micro-CT and fluorescent microphotography to investigate the exceptionally preserved ventral morphology of R. abnormalis and explore its phylogenetic implications through maximum parsimony and Bayesian inference. Detailed morphology revealed here better supports R. abnormalis as a sister group to the diminutive artiopod Pygmaclypeatus daziensis, also known from Chengjiang, and strengthens the close relationship of these taxa that have been suggested by previous studies as early-branching representatives of Trilobitomorpha. Cephalic appendages suggest this animal might be a scavenger, possibly feeding on soft-bodied organisms. Different pairs of pygidial appendages suggest an anamorphic post-embryonic ontogeny, which adds to the understanding of the developmental mode of Cambrian artiopods, and further supports the statement that post-hatching segment addition occurred in the ancestor of Euarthropoda.

Before trilobite legs: Pygmaclypeatus daziensis reconsidered and the ancestral appendicular organization of Cambrian artiopods.

The Cambrian Stage 3 Chengjiang biota in South China is one of the most influential Konservat-Lagerstätten worldwide thanks to the fossilization of diverse non-biomineralizing organisms through pyritization. Despite their contributions to understanding the evolution of early animals, several Chengjiang species remain poorly known owing to their scarcity and/or incomplete preservation. Here, we use micro-computed tomography to reveal in detail the ventral appendage organization of the enigmatic non-trilobite artiopod Pygmaclypeatus daziensis-one of the rarest euarthropods in Chengjiang-and explore its functional ecology and broader evolutionary significance. Pygmaclypeatus daziensis possesses a set of uniramous antennae and 14 pairs of post-antennal biramous appendages, the latter of which show an unexpectedly high degree of heteronomy based on the localized differentiation of the protopodite, endopodite and exopodite along with the antero-posterior body axis. The small body size (less than 2 cm), the presence of delicate spinose endites and well-developed exopodites with multiple paddle-shaped lamellae on the appendages of P. daziensis indicate a nekto-benthic mode of life and a scavenging/detritus feeding strategy. Pygmaclypeatus daziensis shows that appendage heteronomy is phylogenetically widespread within Artiopoda-the megadiverse clade that includes trilobites and their relatives with non-biomineralizing exoskeletons-and suggests that a single exopodite lobe with paddle-like lamellae is ancestral for this clade. This article is part of the theme issue 'The impact of Chinese palaeontology on evolutionary research'.

Spines and baskets in apex predatory sea scorpions uncover unique feeding strategies using 3D-kinematics.

Megalograptidae and Mixopteridae with elongate, spinose prosomal appendages are unique early Palaeozoic sea scorpions (Eurypterida). These features were presumably used for hunting, an untested hypothesis. Here, we present 3D model-based kinematic range of motion (ROM) analyses of Megalograptus ohioensis and Mixopterus kiaeri and compare these to modern analogs. This comparison confirms that the eurypterid appendages were likely raptorial, used in grabbing and holding prey for consumption. The Megalograptus ohioensis model illustrates notable Appendage III flexibility, indicating hypertrophied spines on Appendage III may have held prey, while Appendage II likely ripped immobilized prey. Mixopterus kiaeri, conversely, constructed a capture basket with Appendage III, and impaled prey with Appendage II elongated spines. Thus, megalograptid and mixopterid frontalmost appendages constructed a double basket system prior to moving dismembered prey to the chelicerae. Such 3D kinematic modeling presents a more complete understanding of these peculiar euchelicerates and highlights their possible position within past ecosystems.

Kinematics of whip spider pedipalps: a 3D comparative morpho-functional approach.

Amblypygi are tropical and subtropical ambush predators that use elongated, raptorial pedipalps for different activities. Although pedipalp use in predation and courtship has been explored in videography in vivo analyses, kinematic ex vivo examination of these appendages has not been conducted. Here, we rectify this lack of data by using micro-CT scans to 3D-kinematically model the appendage morphology and the range of motion (ROM) of the joints for Damon medius and Heterophrynus elaphus. We illustrate the successful application of this technique to terrestrial euarthropods in determining the maximum ROM values for each pedipalp joint. We also note that, in life, these values would be lower due to motion restricting structures like tendons, arthrodial membranes, and muscles. We further compare our maximum values obtained here with data from video-based motion analyses. The ROM of each joint shows the greatest flexibility in the femur-tibia joint (140-150°), the lowest in the basitarsus-claw joint (35-40°). ROM in the tibia-basitarsus joint is markedly distinct (D. medius: 44°; H. elaphus: 105°). This disparity reflects how H. elaphus uses the joint in the capture basket, while D. medius uses the femur-tibia joint to form the capture basket. We further illustrate notable vertical motion of the H. elaphus pedipalp compared to D. medius. This difference reflects the retro-ventral trochanter apophysis of H. elaphus. Our study opens the possibility to further whip spider kinematic understanding. Examination of other taxa using this approach will result in a more comprehensive understanding of the ecological significance and ethological implications of this unique arachnid group.

Exites in Cambrian arthropods and homology of arthropod limb branches.

The last common ancestor of all living arthropods had biramous postantennal appendages, with an endopodite and exopodite branching off the limb base. Morphological evidence for homology of these rami between crustaceans and chelicerates has, however, been challenged by data from clonal composition and from knockout of leg patterning genes. Cambrian arthropod fossils have been cited as providing support for competing hypotheses about biramy but have shed little light on additional lateral outgrowths, known as exites. Here we draw on microtomographic imaging of the Cambrian great-appendage arthropod Leanchoilia to reveal a previously undetected exite at the base of most appendages, composed of overlapping lamellae. A morphologically similar, and we infer homologous, exite is documented in the same position in members of the trilobite-allied Artiopoda. This early Cambrian exite morphology supplements an emerging picture from gene expression that exites may have a deeper origin in arthropod phylogeny than has been appreciated.

Intraspecific variation in the Cambrian: new observations on the morphology of the Chengjiang euarthropod Sinoburius lunaris.

BACKGROUND: The Chengjiang biota from southwest China (518-million-years old, early Cambrian) has yielded nearly 300 species, of which more than 80 species represent early chelicerates, crustaceans and relatives. The application of µCT-techniques combined with 3D software (e.g., Drishti), has been shown to be a powerful tool in revealing and analyzing 3D features of the Chengjiang euarthropods. In order to address several open questions that remained from previous studies on the morphology of the xandarellid euarthropod Sinoburius lunaris, we reinvestigated the µCT data with Amira to obtain a different approach of visualization and to generate new volume-rendered models. Furthermore, we used Blender to design 3D models showing aspects of intraspecific variation. RESULTS: New findings are: (1) antennulae consist of additional proximal articles that have not been detected before; (2) compared to other appendages, the second post-antennular appendage has a unique shape, and its endopod is comprised of only five articles (instead of seven); (3) the pygidium bears four pairs of appendages which are observed in all specimens. On the other hand, differences between specimens also have been detected. These include the presence/absence of diplotergites resulting in different numbers of post-antennular appendages and tergites and different distances between the tip of the hypostome and the anterior margin of the head shield. CONCLUSIONS: Those new observations reveal intraspecific variation among Chengjiang euarthropods not observed before and encourage considerations about possible sexual dimorphic pairs or ontogenetic stages. Sinoburius lunaris is a variable species with respect to its morphological characters, cautioning that taxon-specific variabilities need to be considered when exploring new species.

Moving legs: A workflow on how to generate a flexible endopod of the 518 million-year-old Chengjiang arthropod Ercaicunia multinodosa using 3D-kinematics (Cambrian, China).

Understanding the functional morphology and mobility of appendages of fossil animals is important for exploring ecological traits such as feeding and locomotion. Previous work on fossils from the 518 million-year-old Chengjiang biota of China was based mainly on two-dimensional information captured from the surface of the specimens. Only recently, µCT techniques started to reveal almost the entire, though flattened and compressed, three-dimensionally preserved morphologies of the arthropods from Chengjiang. This allows more accurate work on reconstructing the possible movement of certain structures such as the appendages. Here, we present a workflow on how to reconstruct the mobility of a limb of the early Chengjiang arthropod Ercaicunia multinodosa from the famous Chinese fossil site. Based on µCT scans of the fossil, we rendered surface models of the 13th-15th right endopods using the 3D visualization and 3D-rendering software Amira. The 3D objects then were postprocessed (Collapse Hierarchy, Unify Normals) in SAP 3D Visual Enterprise Author before being imported into the 3D animation program Autodesk Maya 2020. Using the add-on tool X_ROMM in Maya, we illustrate step-by-step on how to make the articles of the limbs swing-in toward each other. Eventually, we propose several possible limb movements of E. multinodosa, which helps to understand how this early arthropod could have moved its endopods.

Outsourcing a visual neuropil - The central visual system of the median eyes of Galeodes granti Pocock, 1903 (Arachnida: Solifugae).

Only a few studies have examined the central visual system of Solifugae until now. To get new insights suitable for phylogenetic analysis we studied the R-cell (or retinula cell) projections and visual neuropils of Galeodes granti using various methods. G. granti possesses large median eyes and rudimentary lateral eyes. In this study, only the R-cells and neuropils of the median eyes were successfully stained. The R-cells terminate in two distinct visual neuropils. The first neuropil is located externally to the protocerebrum directly below the retina, the second neuropil lies in the cell body rind of the protocerebrum, and immediately adjacent is the arcuate body. This layout of the median eye visual system differs from Arachnopulmonata (Scorpiones + Tetrapulmonata). However, there are several similarities with Opiliones. In both, (1) the R-cells are connected to a first and second visual neuropil and not to any other region of the brain, (2) the first neuropil is not embedded in the cell body rind of the protocerebrum, it is rather external to the protocerebrum, (3) the second visual neuropil is embedded in the cell body rind, and (4) the second neuropil abuts the arcuate body. These findings may provide important new characters for the discussion on arachnid phylogeny.

Auditory chain reaction: Effects of sound pressure and particle motion on auditory structures in fishes.

Despite the diversity in fish auditory structures, it remains elusive how otolith morphology and swim bladder-inner ear (= otophysic) connections affect otolith motion and inner ear stimulation. A recent study visualized sound-induced otolith motion; but tank acoustics revealed a complex mixture of sound pressure and particle motion. To separate sound pressure and sound-induced particle motion, we constructed a transparent standing wave tube-like tank equipped with an inertial shaker at each end while using X-ray phase contrast imaging. Driving the shakers in phase resulted in maximised sound pressure at the tank centre, whereas particle motion was maximised when shakers were driven out of phase (180°). We studied the effects of two types of otophysic connections-i.e. the Weberian apparatus (Carassius auratus) and anterior swim bladder extensions contacting the inner ears (Etroplus canarensis)-on otolith motion when fish were subjected to a 200 Hz stimulus. Saccular otolith motion was more pronounced when the swim bladder walls oscillated under the maximised sound pressure condition. The otolith motion patterns mainly matched the orientation patterns of ciliary bundles on the sensory epithelia. Our setup enabled the characterization of the interplay between the auditory structures and provided first experimental evidence of how different types of otophysic connections affect otolith motion.

Combining morphological and genomic evidence to resolve species diversity and study speciation processes of the Pallenopsis patagonica (Pycnogonida) species complex.

BACKGROUND: Pallenopsis patagonica (Hoek, 1881) is a morphologically and genetically variable sea spider species whose taxonomic classification is challenging. Currently, it is considered as a species complex including several genetic lineages, many of which have not been formally described as species. Members of this species complex occur on the Patagonian and Antarctic continental shelves as well as around sub-Antarctic islands. These habitats have been strongly influenced by historical large-scale glaciations and previous studies suggested that communities were limited to very few refugia during glacial maxima. Therefore, allopatric speciation in these independent refugia is regarded as a common mechanism leading to high biodiversity of marine benthic taxa in the high-latitude Southern Hemisphere. However, other mechanisms such as ecological speciation have rarely been considered or tested. Therefore, we conducted an integrative morphological and genetic study on the P. patagonica species complex to i) resolve species diversity using a target hybrid enrichment approach to obtain multiple genomic markers, ii) find morphological characters and analyze morphometric measurements to distinguish species, and iii) investigate the speciation processes that led to multiple lineages within the species complex. RESULTS: Phylogenomic results support most of the previously reported lineages within the P. patagonica species complex and morphological data show that several lineages are distinct species with diagnostic characters. Two lineages are proposed as new species, P. aulaeturcarum sp. nov. Dömel & Melzer, 2019 and P. obstaculumsuperavit sp. nov. Dömel, 2019, respectively. However, not all lineages could be distinguished morphologically and thus likely represent cryptic species that can only be identified with genetic tools. Further, morphometric data of 135 measurements showed a high amount of variability within and between species without clear support of adaptive divergence in sympatry. CONCLUSIONS: We generated an unprecedented molecular data set for members of the P. patagonica sea spider species complex with a target hybrid enrichment approach, which we combined with extensive morphological and morphometric analyses to investigate the taxonomy, phylogeny and biogeography of this group. The extensive data set enabled us to delineate species boundaries, on the basis of which we formally described two new species. No consistent evidence for positive selection was found, rendering speciation in allopatric glacial refugia as the most likely model of speciation.

The visual system of Thelyphonida (whip scorpions): Support for Arachnopulmonata.

Only a few studies have examined the central visual system of Thelyphonida (whip scorpions) until now. To obtain new insights suitable for phylogenetic analysis we studied the axonal trajectories and neuropil architecture of the central visual systems of two whip scorpion species (Mastigoproctus giganteus and Typopeltis dalyi) with different neuroanatomical techniques (Cobalt fills, Wigglesworth stains, and µCT). The central visual system of whip scorpion comprises one pair of median eyes and one pair of lateral eye triplets. The R-cells (or retinula cells) of both eye types each terminate in a first and a second visual neuropil. Furthermore, a few R-cell fibres from the median eyes leave the second median eye visual neuropil and terminate in the second and the first lateral eye neuropil. This means R-cell terminals from the lateral eyes and the median eyes overlap here. Additionally, the arcuate body and mushroom bodies are described. A detailed comparison of our findings with previously studied chelicerate central visual systems seems to support a monophyly of Arachnopulmonata, i.e. a clade comprising Tetrapulmonata (Thelyphonida, Schizomida, Amblypygi, and Araneae) and Scorpions. Furthermore, the architecture of the central visual systems hints at a close evolutionary relationship of Arachnopulmonata and Xiphosura.

Also looking like Limulus? - retinula axons and visual neuropils of Amblypygi (whip spiders).

BACKGROUND: Only a few studies have examined the visual systems of Amblypygi (whip spiders) until now. To get new insights suitable for phylogenetic analysis we studied the axonal trajectories and neuropil architecture of the visual systems of several whip spider species (Heterophrynus elaphus, Damon medius, Phrynus pseudoparvulus, and P. marginemaculatus) with different neuroanatomical techniques. The R-cell axon terminals were identified with Cobalt fills. To describe the morphology of the visual neuropils and of the protocerebrum generally we used Wigglesworth stains and µCT. RESULTS: The visual system of whip spiders comprises one pair of median and three pairs of lateral eyes. The R-cells of both eye types terminate each in a first and a second visual neuropil. Furthermore, a few R-cell fibres from the median eyes leave the second median eye visual neuropil and terminate in the second lateral eye neuropil. This means R-cell terminals from the lateral eyes and the median eyes overlap. Additionally, the arcuate body and the mushroom bodies are described. CONCLUSIONS: A detailed comparison of our findings with previously studied chelicerate visual systems (i.e., Xiphosura, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae) seem to support the idea of close evolutionary relationships between Xiphosura, Scorpiones, and Amblypygi.

Feeding ecology in sea spiders (Arthropoda: Pycnogonida): what do we know?

Sea spiders (Pycnogonida) are a widespread and phylogenetically important group of marine arthropods. However, their biology remains understudied, and detailed information about their feeding ecology is difficult to find. Observations on pycnogonid feeding are scattered in the literature, often in older sources written in various languages, and have never been comprehensively summarized. Here we provide an overview of all information on feeding in pycnogonids that we have been able to find and review what is known on feeding specializations and preferences in the various pycnogonid taxa. We deduce general findings where possible and outline future steps necessary to gain a better understanding of the feeding ecology of one of the world's most bizarre animal taxa.

In-situ visualization of sound-induced otolith motion using hard X-ray phase contrast imaging.

Regarding the basics of ear structure-function relationships in fish, the actual motion of the solid otolith relative to the underlying sensory epithelium has rarely been investigated. Otolith motion has been characterized based on a few experimental studies and on approaches using mathematical modeling, which have yielded partially conflicting results. Those studies either predicted a simple back-and-forth motion of the otolith or a shape-dependent, more complex motion. Our study was designed to develop and test a new set-up to generate experimental data on fish otolith motion in-situ. Investigating the basic parameters of otolith motion requires an approach with high spatial and temporal resolution. We therefore used hard X-ray phase contrast imaging (XPCI). We compared two anatomically well-studied cichlid species, Steatocranus tinanti and Etroplus maculatus, which, among other features, differ in the 3D shape of their otoliths. In a water-filled tank, we presented a pure tone of 200 Hz to 1) isolated otoliths embedded in agarose serving as a simple model or 2) to a fish (otoliths in-situ). Our new set-up successfully visualized the motion of otoliths in-situ and therefore paves the way for future studies evaluating the principles of otolith motion.

The visual system of harvestmen (Opiliones, Arachnida, Chelicerata) - a re-examination.

BACKGROUND: The visual systems in chelicerates are poorly understood, even though they show strong variation in eye and visual neuropil architecture, thus may provide valuable insights for the understanding of chelicerate phylogeny and eye evolution. Comparable morphological characters are desperately sought for reconstructions of the phylogeny of Chelicerata, especially with respect to Arachnida. So far, reliable data exist only for Pycnogonida, Xiphosura, Scorpiones, and Araneae. The few earlier studies of the organisation of the visual system in harvestmen are contradictory concerning the number, morphology, and position of the visual neuropils. RESULTS: We undertook a descriptive and comparative analysis of the neuroanatomy of the visual system in several phalangid harvestmen species. Various traditional and modern methods were used that allow comparisons with previous results (cobalt fills, DiI/DiO labelling, osmium ethyl gallate procedure, and TEM). The R-cells (photoreceptor and arhabdomeric cells) in the eyes of Opiliones are linked to a first and a second visual neuropil. The first visual neuropil receives input from all R-cell axons, in the second only few R-cells terminate in the distal part. Hence, the second visual neuropil is subdivided in a part with direct R-cell input and a part without. The arcuate body is located in a subsequent position with direct contact to the second visual neuropil. CONCLUSIONS: This re-examination comes to conclusions different from those of all previous studies. The visual system of phalangid Opiliones occupies an intermediate position between Pycnogonida, Xiphosura, and Scorpiones on the one side, and Araneae on the other side. The projection of the R-cells is similar to that in the former grouping, the general neuropil arrangement to that in the latter taxon. However, more research on the visual systems in other chelicerate orders is needed in order to draw inferences on phylogeny or eye evolution.

Three-dimensionally preserved minute larva of a great-appendage arthropod from the early Cambrian Chengjiang biota.

A three-dimensionally preserved 2-mm-long larva of the arthropod Leanchoilia illecebrosa from the 520-million-year-old early Cambrian Chengjiang biota of China represents the first evidence, to our knowledge, of such an early developmental stage in a short-great-appendage (SGA) arthropod. The larva possesses a pair of three-fingered great appendages, a hypostome, and four pairs of well-developed biramous appendages. More posteriorly, a series of rudimentary limb Anlagen revealed by X-ray microcomputed tomography shows a gradient of decreasing differentiation toward the rear. This, and postembryonic segment addition at the putative growth zone, are features of late-stage metanauplii of eucrustaceans. L. illecebrosa and other SGA arthropods, however, are considered representative of early chelicerates or part of the stem lineage of all euarthropods. The larva of an early Cambrian SGA arthropod with a small number of anterior segments and their respective appendages suggests that posthatching segment addition occurred in the ancestor of Euarthropoda.