A treat of Cambrian arthropods

Walossek, D. 1993: The Upper Cambrian Rehbachiella and the phylogeny of Branchiopoda and Crustacea. Fossils and Strata 32 .     

Theory of microtomy artefacts in arthropod cuticle

Arthropod cuticles observed in section generally present alternating clear and dark bands. These have often been interpreted in terms of superimposed layers of different structure or composition. It has been shown, however that this material is homogeneous, and is formed by a twisted arrangement of microfibrils. The dense bands correspond actually to a microtomy artefact and they form dark single spirals in certain distorted areas of the cuticle. A model was proposed, involving the interaction between knife motion and microfibrils; it will be referred to as the stepped model, since the proposed mechanism results in the formation of steps at the surface of sections, on both faces. These steps are limited by structures resembling crests or cliffs, whose regular distribution produces alternating thick and thin bands in the section. This explains the observed contrast (Bouligand, (1972)). Two very interesting models were proposed later (Gordon and Winfree, (1978)) and are referred to as the cos F model and the sand model, but steps and crests are absent in these models. However, Giraud-Guille ((1986)), has shown very clearly the existence of these crests, which seem to be quite essential in this microtomy artefact. To clarify the debate, the texts defining the initial and the two new models are reproduced here and the difficulties encountered by each model are discussed. A mathematical formulation of this artefact is presented in an appendix to the present article; this leads to a more complete discussion of the possible models. Other factors are also taken into consideration: microfibril orientation and staining. The main factor of contrasts is undoubtedly the variation in thickness over a single section, as proposed in the stepped model.     

Microfiber reinforcement of an arthropod cuticle

Summary 1. According to its fine structure arthropod cuticle is a laminated composite material. The exoskeleton of a spider is surveyed in an electron microscopical study as to the characteristics of its fiber reinforcement.2. Five types of fiber arrangement are distinguished, ranging from unidirectional to continuously rotating fiber orientation and with three types combining elements of both of these extreme cases. The unidirectional type is rare and its occurrence confined to the innermost part of the walking leg endocuticle and to parts of articular membranes. Quite contrary a continuous and regular rotation of fiber direction is common. In hard cuticle (exocuticle) this type of reinforcement is found exclusively.3. Lamellar width, indicating a 180° rotation of the fiber direction, varies within wide limits. Variation is due to varying numbers of fiber layers and the amount of directional change between them (ca. 0.6–24°, type E).4. The diameter of the chitin/protein microfiber amounts to ca. 35 Å. Its length measured up to 1,5 m in the preparations. A periodicity of ca. 80 Å is noted. The precision of fiber alignment within the cuticle is high.5. In the cuticle of the opisthosoma and the articular membranes endocuticular lamellae are taking a wavy course, which is interpreted to be in relation to the cuticle's softness.6. The mechanical implications of both an increasing number of fiber directions in a composite material and of the different types of fiber reinforcement found in the spider cuticle are demonstrated by a calculation of the directional properties of the moduli of elasticity in tension and shear. The predictable strong influence of both the fiber content and the binder material's E-modulus on the mechanical properties of cuticle is illustrated.7. Instead of a trajectorial design a continuous rotation of the fiber orientation is found in all exocuticles. Thus the spider does not make use of the special advantage (high strength/weight ratio) of unidirectional fiber arrangement in places where it seemed most likely.Many thanks are due to Ms. H. Tscharntke for expert assistance with electron microscopy and photographic work. Dipl.-Ing. F. Och and Dr. G. Haberl (Messerschmitt-Bölkow-Blohm GmbH, Ottobrunn) provided the calculations for Figs. 14 and 15. Their generous help is gratefully acknowledged. Dr. K. Frahm (Universität München, Fakultät f. Tiermedizin) kindly shared his densitometer. Ms. F. Althaus carefully carried out the drawings.     

Trace fossils of Cambrian aglaspidid arthropods

A new trace fossil, Raaschichnus gundersoni , from the Upper Cambrian St. Lawrence Formation of Wisconsin, was produced by an aglaspidid arthropod. The rusophyciform trace, which occurs singly and in series, is distinctive in showing marks left by the tail-spine. Other trace fossils previously considered to have been made by aglaspidids were probably excavated by animals lacking a tail-spine and with appendage morphologies very different from the aglaspidids of Wisconsin.     

Digestive system and feeding mode in Cambrian naraoiid arthropods

The function of the digestive system of naraoiid arthropods is interpreted in the light of new observations on Early Cambrian specimens from China and detailed comparisons with Recent crustaceans and other arthropods. In naraoiids, paired tubular diverticulae ending as blind caeca are present along the entire midgut, and are interpreted as sites for the secretion of digestive enzymes. Naraoia bears one pair of long, ramifying, distensible diverticulae, possibly used for both food storage and digestion as suggested by Recent analogues (e.g. branchiuran and isopod crustaceans and limulids). Naraoiids were probably epibenthic scavengers/predators rather than mud-eaters. They were either opportunistic intermittent feeders ( Naraoia ) or more regular feeders ( Misszhouia ). The mud-fills of the alimentary canals are likely to be artefacts due to taphonomic and weathering processes or, less likely, to sediment ingestion by animals trapped alive in turbiditic flows. The case study of naraoiid arthropods adds to other fossil evidence supporting the idea that predation played a key role in the Early Cambrian food-webs and that organs adapted for this purpose had already reached a high level of diversity and anatomical sophistication.     

The cuticle of the aglaspidid arthropods, a red-herring in the early history of the vertebrates

The supposed early vertebrate Anatolepis has recently been reinterpreted by Peel (1979; Rapp. Gr∅nl. Geol. Unders. 91 ) as an arthropod, probably aglaspidid. The cuticle of Aglaspis shows a characteristic arthropodan structure, including laminations and ducts, but it is apparently composed of primary phosphate. Apart from composition, the skeletal histology of Anatolepis is totally different, and this, together with more general considerations, eliminates any possibility of arthropod affinity. Anatolepis remains the earliest known vertebrate. Aglaspididae, cuticle, ultrastructure, Vertebrata, Upper Cambrian.     

Cambrian bivalved arthropod reveals origin of arthrodization

Extant arthropods are diverse and ubiquitous, forming a major constituent of most modern ecosystems. Evidence from early Palaeozoic Konservat Lagerstätten indicates that this has been the case since the Cambrian. Despite this, the details of arthropod origins remain obscure, although most hypotheses regard the first arthropods as benthic predators or scavengers such as the fuxianhuiids or megacheirans (‘great-appendage’ arthropods). Here, we describe a new arthropod from the Tulip Beds locality of the Burgess Shale Formation (Cambrian, series 3, stage 5) that possesses a weakly sclerotized thorax with filamentous appendages, encased in a bivalved carapace, and a strongly sclerotized, elongate abdomen and telson. A cladistic analysis resolved this taxon as the basal-most member of a paraphyletic grade of nekto-benthic forms with bivalved carapaces. This grade occurs at the base of Arthropoda (panarthropods with arthropodized trunk limbs) and suggests that arthrodization (sclerotization and jointing of the exoskeleton) evolved to facilitate swimming. Predatory and fully benthic habits evolved later in the euarthropod stem-lineage and are plesiomorphically retained in pycnogonids (sea spiders) and euchelicerates (horseshoe crabs and arachnids).     

Cephalic and appendage morphology of the Cambrian arthropod Sidneyia inexpectans

Sidneyia inexpectans Walcott, 1911 from the Cambrian Series 3 Burgess Shale of British Columbia is largely accepted as a representative of the artiopodans, an assemblage of Paleozoic arthropod taxa, including trilobites and their immediate relatives. Its appendage morphology was never fully understood, but the exopod seemed to differ from that of other artiopodans, except for the shared presence of lamellae. The head was considered to comprise only the ocular and antennular segments, these being covered entirely on the ventral side by a large doublure. This short head was often taken as an evidence for variability of head segment counts in Cambrian arthropods, and to falsify the hypothesis of a head with three postantennular segments in the euarthropod ground pattern. Restudy of a substantial amount of material of S. inexpectans shows that previous interpretations of a short head were based on taphonomically deformed specimens, where the head was either partly folded, or entirely flipped under the thorax, resulting in the dorsal shield being mistaken for an extensive doublure. Rather than an extensive doublure, there is a broad hypostome, and the head comprises ocular, antennular, and at least two postantennular appendage bearing segments. The appendage morphology is shown to be consistent with artiopodan affinities. The exopod is of the bilobate flap-like type with lamellae inserting on the proximal portion, earlier proposed as a potential autapomorphy of Artiopoda. Reinforcement of artiopodan affinities for S. inexpectans and reinterpretation of its head reconciles this species with current understanding of arthropod phylogeny and head segmentation.     

The nature of driving forces for passive water transport through arthropod cuticle

1. 1.|We demonstrate using thermodynamic arguments that water loss through arthropod epicuticle is well described by a linear law relating water flux to transmembrane vapour pressure drop.

2. 2.|The relationship applies equally to systems where the liquid or vapour exist on either side of a membrane.

3. 3.|An earlier claim by some workers that water diffusion through arthropod epicuticle is proportional to chemical potential drop across the membrane is found to be theoretically unjustified.

4. 4.|Recent measurements with Periplaneta cuticle support the prediction that flux at a given temperature is proportional to the difference in vapour pressure.

A Parvancorina-like arthropod from the Cambrian of South China

Constraining the origin of animal groups is allowed, to some extent, by discoveries of Cambrian Lagerstätten that preserve both mineralizing and nonmineralizing organisms. A new species is reported here of the Cambrian arthropod Skania, which bears an exoskeleton that shares homologies with the Neoproterozoic (Ediacaran) organism Parvancorina and firmly establishes a Precambrian root for arthropods. A new monophyletic group, Parvancorinomorpha, is proposed as the first clade within the arthropod crown group demonstrably ranging across the Neoproterozoic–Paleozoic transition. The Parvancorinomorpha is interpreted to be the sister group of the Arachnomorpha. Incipient cephalization in Skania and related genera represents a step in the progression toward division of a cephalon from a large posterior trunk as shown in Cambrian arachnomorphs such as naraoiids and the addition of a pygidium and thoracic tergites as shown in the arachnomorph clade basal to trilobites. This evidence can serve as a new calibration point for estimating the divergence time for the last common ancestor of arthropods and priapulids based on molecular clock methods.     

The arthropod fauna of Quebec vineyards with particular reference to phytophagous arthropods

A 3-yr study using different sampling and trapping techniques showed that the arthropod pest fauna in two commercial vineyards in southwestern Quebec was qualitatively and quantitatively different than that of Ontario, Canada, and New York state. We hypothesize that a colder winter climate in addition to the agronomic activity of earthing up around the vines in autumn to protect the roots from freezing in winter contributed to low numbers of pests, such as the grape berry moth, Endopiza viteana Clemens (Lepidoptera, Tortricidae). Once in 3 yr, the density of this pest approached, in one of the vineyards, the action threshold recommended for New York. Therefore, it should be monitored on an annual basis. Another phytophagous arthropod that has the potential to cause sporadic economic damage is the potato leafhopper, Empoasca fabae (Harris). The Asiatic garden beetle, Maladera (= Autoserica) castanea (Arrow), was reported for the first time in Canada. The tarnished plant bug, Lygus lineolaris (Palisot de Beauvois), was also captured by sampling. However, its status as a pest has yet to be clarified.     

A myriapod-like arthropod from the Upper Cambrian of East Siberia

Although the fossil record of biramous arthropods commences in the Lower Cambrian, unequivocal uniramous arthropods do not appear until the Upper Silurian, in association with terrestrial biotas. Here we report an Upper Cambrian marine arthropod from East Siberia that possesses some significant myriapodan features. The new arthropod,Xanthomyria spinosa n. gen., n. sp., closely resembles examples of archipolypodans from the Late Palaeozoic. If this resemblance genuinely represents myriapod affinities, this would be the first convincing myriapod from the Cambrian. Suggestions of an early branching point of the myriapods from other arthropods would be consistent with this. Conversely, an as yet poorly known clade of multi-segmented arthropods may exist in the Cambrian, with no close affinities to the myriapods.      

Hydrodynamics and sliding posture analysis of the Cambrian arthropod Ercaicunia multinodosa

Dynamic mechanical analysis offers the opportunity to explore the motility and feeding strategies of extinct organisms, but the prerequisite is to have an accurate recovery engineering model. As shown with micro-computed tomography (CT) scanning, fossils of the bivalved arthropod Ercaicunia multinodosa from the Cambrian (Series 2, Stage 3) Chengjiang biota of China have well-preserved three-dimensional (3D) morphological details with a certain degree of compression. Here, we propose a palaeontological restoration method using computational fluid dynamics (CFD) to analyse multiple hypothetical models based on the fossil information to obtain a reasonable restoration model. Furthermore, we carry out hydrodynamic experiments to verify the palaeontological restoration results. Our simulation and experimental results suggest that, a dorsally convex and ventrally straight body shape with the valves opening at an angle of approximately 120° works best for E. multinodosa to overcome resistance, and in the meantime obtain most lift while sliding in the water column. The combination of three-dimensional reconstruction, CFD simulation, and hydrodynamic experiments provides a useful method for restoring the morphologies of extinct animals and exploring their palaeoecology.