Spinicaudatans and ostracods (Crustacea) from the Montceau Lagerstätte (Late Carboniferous, France): morphology and palaeoenvironmental significance

Spinicaudatans and ostracods form two components of the diverse arthropod fauna from the Montceau Lagerstätte (Stephanian, France). Spinicaudatans are represented by Montcestheria orri gen. and sp. nov. and Euestheria feysi sp. nov., and ostracods by a single species, Carbonita sp. aff. salteriana (Jones, 1862). Allied forms such as Montcestheria sp. aff. orri, Montcestheria sp. and Euestheria cebennensis (Grand'Eury, 1890), all from coeval localities in France, are also described. Montcestheria gen. nov. has carapace features, external (possibly sexual) dimorphism, preserved soft parts (e.g. appendages, gut) and resting eggs similar to those of Recent spinicaudatans, suggesting comparable lifestyles, reproductive strategies and feeding modes. Detailed anatomical comparisons are made with Cyzicus tetracerus from Recent ephemeral freshwater ponds. The ostracods belong to the Superfamily Carbonitoidea, which is a recurrent component of Carboniferous non‐marine biotas. Spinicaudatan‐rich assemblages typically occur in several Westphalian–Stephanian Lagerstätten (including Montceau) and localities from Europe and North America, where freshwater conditions prevailed, indicating that the group had already colonized continental waters by the Late Carboniferous. Similarities with the fauna from Recent temporary freshwater ponds (e.g. low diversity/high density spinicaudatan‐ostracod populations, synchronous spinicaudatan populations developing from resting eggs, high diversity/low density insects, amphibians) suggest the presence of temporary or ephemeral aquatic environments at Montceau as part of a complex limnic ecosystem. Flooding may have been the main driving force by which faunal and floral elements drifted away from their respective biotopes into the depositional areas, thus explaining the co‐occurrence of terrestrial (e.g. myriapods, scorpions, plants), amphibian and aquatic (e.g. conchostracans and syncarids from temporary and permanent settings, respectively) elements in fossil assemblages.     

Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China

The Chengjiang Lagerstätte in the Lower Cambrian of South China yields a small, larva‐like arthropod, which was considered to be a protaspis of naraoiids by many authors. The discovery of a large number of well‐preserved specimens from many new localities has allowed the original study to be revised. The relatively large size, stable morphology and unusual structure of the appendages indicate that these specimens represent adults of a new arthropod, Primicaris larvaformis. The larva‐like outline is considered to have arisen by the heterochronic process of progenesis. In addition, this animal displays primitive aspects of bodyplan and limb morphology that suggest a basal position within arachnomorphs, or perhaps even arthropods, and the similarities to the Vendian arthropod‐like animal Parvancorina probably provide an evolutionary link between Vendian forms and Cambrian arthropods.     

Carboniferous Onychophora from Montceau‐les‐Mines,France, and onychophoran terrestrialization

The geological age of the onychophoran crown‐group, and when the group came onto land, have been sources of debate. Although stem‐group Onychophora have been identified from as early as the Cambrian, the sparse record of terrestrial taxa from before the Cretaceous is subject to contradictory interpretations. A Late Carboniferous species from the Mazon Creek biota of the USA, Helenodora inopinata, originally interpreted as a crown‐group onychophoran, has recently been allied to early Cambrian stem‐group taxa. Here we describe a fossil species from the Late Carboniferous Montceau‐les‐Mines Lagerstätte, France, informally referred to as an onychophoran for more than 30 years. The onychophoran affinities of Antennipatus montceauensis gen. nov., sp. nov. are indicated by the form of the trunk plicae and the shape and spacing of their papillae, details of antennal annuli, and the presence of putative slime papillae. The poor preservation of several key systematic characters for extant Onychophora, however, prohibits the precise placement of the Carboniferous fossil in the stem or crown of the two extant families, or the onychophoran stem‐group as a whole. Nevertheless, A. montceauensis is the most compelling candidate to date for a terrestrial Paleozoic onychophoran.     

A rare non‐trilobite artiopodan from the Guzhangian (Cambrian Series 3) Weeks Formation Konservat‐Lagerstätte in Utah,USA

We describe a weakly biomineralized non‐trilobite artiopodan arthropod from the Guzhangian Weeks Formation of Utah. Falcatamacaris bellua gen. et sp. nov. is typified by a thin calcitic cuticle, broad cephalon without eyes or dorsal ecdysial sutures, an elongate trunk with distinctively sickle‐shaped pleural spines and a long tailspine with a bifurcate termination. The precise affinities of Falcatamacaris gen. nov. are problematic due to the presence of unique features within Artiopoda, such as the peculiar morphology of the pleural and posterior regions of the trunk. Possible affinities with aglaspidid‐like arthropods and concilitergans are discussed based on the possession of 11 trunk tergites, edge‐to‐edge articulations and overall body spinosity. The new taxon highlights the importance of the Weeks Formation Konservat‐Lagerstätte for further understanding the diversity of extinct arthropod groups in the upper Cambrian.     

Morpho-anatomy of the lobopod Magadictyon cf. haikouensis from the Early Cambrian Chengjiang Lagerstätte, South China

Magadictyon haikouensis (Luo and Hu, 1999) from the Early Cambrian Chengjiang Lagerstätte, an incomplete specimen of a large lobopod with strong appendages, has been regarded as related to the lobopods Microdictyon and Onychodictyon. Newly discovered complete specimens of Magadictyon cf. haikouensis (found by the Early Life Institute field team) show that the taxon, in addition to its strong appendages with appendicules, also had a head bearing similar caecum‐like structures to those of the arthropod Naraoia and Chelicerate, ‘Peytoia’‐like mouthparts and frontal appendages. Because of their similarity, the caecum‐like structures of Magadictyon cf. haikouensis are considered to be homologous with those of stem‐group arthropods. The ‘Peytoia’‐like mouthparts and the frontal appendages are similar to those of the AOPK (Anomalocaris–Opabinia–Pambdelurion–Kerygmachela) group. In addition, the appendages with appendicules show that Magadictyon cf. haikouensis is closely related to Onychodictyon. Therefore, Magadictyon cf. haikouensis is regarded here as a rare transitional form between lobopods and arthropods. Besides, together with other lobopods, the morphology of Magadictyon cf. haikouensis demonstrates that the Cambrian lobopods appear to have been diverse and not particularly closely related to one another, and do not seem to represent a monophyletic clade.     

Onychophoran‐like myoanatomy of the Cambrian gilled lobopodian Pambdelurion whittingtoni

Arthropods are characterized by a rigid, articulating, exoskeleton operated by a lever‐like system of segmentally arranged, antagonistic muscles. This skeletomuscular system evolved from an unsegmented body wall musculature acting on a hydrostatic skeleton, similar to that of the arthropods’ close relatives, the soft‐bodied onychophorans. Unfortunately, fossil evidence documenting this transition is scarce. Exceptionally‐preserved panarthropods from the Cambrian Lagerstätte of Sirius Passet, Greenland, including the soft‐bodied stem‐arthropod Pambdelurion whittingtoni and the hard‐bodied arthropods Kiisortoqia soperi and Campanamuta mantonae, are unique in preserving extensive musculature. Here we show that Pambdelurion's myoanatomy conforms closely to that of extant onychophorans, with unsegmented dorsal, ventral and longitudinal muscle groups in the trunk, and extrinsic and intrinsic muscles controlling the legs. Pambdelurion also possesses oblique musculature, which has previously been interpreted as an arthropodan characteristic. However, this oblique musculature appears to be confined to the cephalic region and first few body segments, and does not represent a shift towards arthropodan myoanatomy. The Sirius Passet arthropods, Kiisortoqia and Campanamuta, also possess large longitudinal muscles in the trunk, although, unlike Pambdelurion, they are segmentally divided at the tergal boundaries. Thus, the transition towards an arthropodan myoanatomy from a lobopodian ancestor probably involved the division of the peripheral longitudinal muscle into segmented units.     

New trace fossil evidence for eurypterid swimming behaviour

We describe a recently discovered trace fossil from a eurypterid Konservat‐Lagerstätte in the upper Silurian Tonoloway Formation of Pennsylvania, and formally describe contemporaneous traces from the Williamsville Formation Lagerstätte of Ontario. The traces from both localities are assigned here to Arcuites bertiensis igen. et isp. nov. Based on comparisons with previously described eurypterid trackways, neoichnological experiments, and the co‐occurrence with eurypterid remains, Arcuites is interpreted as having been made by the swimming leg (sixth prosomal appendage) of swimming juvenile to adult eurypteroid eurypterids, and represents the first unambiguous trace fossil evidence for eurypterid swimming behaviour. The morphology of Arcuites indicates that eurypteroid eurypterids swam using drag‐based rowing, whereby the animal propelled itself forward by moving its oar blade‐like swimming paddles in an in‐phase backstroke. Arcuites morphology also indicates that the eurypteroid swimming appendage had a greater degree of movement than was previously suggested, and a revised rowing model is proposed. Differences in the abundance of A. bertiensis in the Tonoloway and Williamsville formations suggest a bathymetric control on eurypterid swimming behaviour and trace production. The association of Arcuites with eurypterid body fossils in both units indicates that these Lagerstätten were autochthonous assemblages and provides additional evidence for eurypterid inhabitation of shallow subtidal marine environments in the late Silurian.     

A new arthropod from the Early Cambrian of North Greenland,with a ‘great appendage’‐like antennula

Kiisortoqia soperi gen. et sp. nov. is an arthropod species from the Early Cambrian Sirius Passet Lagerstätte of North Greenland. A head, incorporating four appendiferous segments and biramous limbs, with an anteroposteriorly compressed basipod with a spine bearing median edge, support the euarthropod affinities of K. soperi gen. et sp. nov. Similarities with ‘short great appendage’ arthropods, or megacheirans, like the nine‐segmented endopod, and the flap‐ or paddle‐like exopod, may be symplesiomorphies. The antennula, however, resembles in composition and size the anteroventral raptorial appendage of anomalocaridids. Thus, the morphology of K. soperi gen. et sp. nov. provides additional support for the homologization of the anomalocaridid ‘great appendage’ with the appendage of the antennular or deutocerebral segment of extant Euarthropoda. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 477–500.     

Probing the deep shelf: a Lagerstätte from the Upper Ordovician of Girvan, southwestern Scotland

A deep‐water Konservat Lagerstätte from the lower Caradoc (Sandbian) at Girvan is dominated by the trilobite Diacanthaspis trippi, the carpoids Anatifopsis n. sp.? and a new genus of ctenocystoid together with the polyplacophoran Solenocaris solenoides and the brachiopod Onniella williamsi. Most of these are multi‐element organisms, with many specimens preserved in an articulated state in finely laminated rocks, indicating minimal disturbance and suggesting that the fauna is largely an in situ association. It contains few of the species known from other deep‐water sites of similar age at Girvan which contain diverse assemblages of trilobites and brachiopods absent from the Lagerstätte. The taphonomy of the site indicates preservation by rapid burial followed by early diagenesis under dysaerobic conditions. It provides a ‘taphonomic window’ on otherwise unknown faunas from distal shelf facies on the Ordovician Laurentian margin, and, moreover, is an important reminder of the hidden biodiversity that resided in thin‐shelled, multi‐element organisms.     

The bivalved arthropods Isoxys and Tuzoia with soft‐part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island,Australia)

Abstract: Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft‐bodied features previously unknown from this Burgess Shale‐type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better‐known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.     

Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschelkalk) mud flat

Knaust, D. 2010: Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschelkalk) mud flat. Lethaia, Vol. 43, pp. 344–356. A new Fossil‐Lagerstätte is reported from the Middle Triassic of Germany, preserving Foraminifera, Nematoda, Platyhelminthes, Nemertea, Annelida and a range of Arthropoda together with their traces. This is the oldest fossil record of free‐living nematodes and turbellarians, and the first occurrence of nemerteans in the Mesozoic. The rare preservation of the benthic associations together with their traces is unique; especially the abundant occurrence of different phyla as meiofauna (organisms with shortest dimension between 0.06 and 1 mm) provides an insight into the palaeoecological conditions of a 240‐Ma‐old muddy tidal flat. The preservation of benthic animals at the termination of their traces offers the exclusive opportunity to assign producers to the trace fossils. The results show that the discussed phyla were already established in the early Mesozoic in a similar diversity and composition as in modern analogues. The new Fossil‐Lagerstätte has the potential to prove a number of soft‐bodied taxa immediately after the end‐Permian mass extinction, which has wide‐ranging implications for phylogenetic interpretations. □Benthos, soft‐bodied organisms, Fossil‐Lagerstätte, Muschelkalk, trace fossils, Triassic.     

Feeding behaviour of a new worm (Priapulida) from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia)

Singuuriqia simoni gen. et sp. nov. represents the first record of a priapulid worm from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia). It is defined by an unusually broad, longitudinally folded, foregut which tapers through the pharynx towards the anterior mouth; posteriorly, the same longitudinal folding is evident in the narrow gut. The slender, smooth, trunk in the unique specimen passes anteriorly into an oval proboscis which culminates in a smooth, extensible, pharynx with pharyngeal teeth. The capacity for substantial expansion of the foregut permitted rapid ingestion of food prior to digestion at leisure. Cololites suggest both carnivorous and deposit feeding behaviour, indicating that Singuuriqia, like the present day Priapulus, was probably omnivorous.     

A new xandarellid arthropod from the Chengjiang Lagerstätte,Lower Cambrian of Southwest China

A new arthropod, Luohuilinella rarus nov. gen., nov. sp., is described from the Lower Cambrian Chengjiang Lagerstätte (Series 2, Stage 3), Yunnan, Southwest China. Luohuilinella nov. gen. is extremely rare in this Lagerstätte, represented by a single specimen. It has a large cephalic shield and a tapering trunk with well-developed pleural field. The cephalic shield is crescentic in outline and its anterolateral margin has two notches. The trunk is weakly trilobate, composed of 27 distinct tergites with well-developed pleural spines and a terminal piece. Luohuilinella nov. gen. resembles xandarellids in overall body architecture and especially in the reduced first trunk tergite. Its ventral morphology is, however, presently unknown. Therefore, it is provisionally assigned to Xandarellida.     

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

Functional morphology,ontogeny and evolution of mantis shrimp‐like predators in the Cambrian

Abstract: We redescribe the morphology of Yohoia tenuis (Chelicerata sensu lato) from the Cambrian Burgess Shale Lagerstätte. The morphology of the most anterior, prominent, so‐called great appendage changes throughout ontogeny. While its principal morphology remains unaltered, the length ratios of certain parts of the great appendage change significantly. Furthermore, it possesses a special jack‐knifing mechanism, i.e. an elbow joint: the articulation between the distal one of the two peduncle elements and the most proximal of the four spine‐bearing claw elements. This morphology might have enabled the animal to hunt like a modern spearer‐type mantis shrimp, an analogy enhanced by the similarly large and protruding eyes. For comparison, details of specimens of selected other great‐appendage arthropods from the Lower Cambrian Chengjiang Lagerstätte have been investigated using fluorescence microscopy. This revealed that the morphology of the great appendage of Y. tenuis is much like that of the Chengjiang species Fortiforceps foliosa and Jianfengia multisegmentalis. The morphology of the great appendage of the latter is even more similar to the morphology developed in early developmental stages of Y. tenuis, while the morphology of the great appendage of F. foliosa is more similar to that of later developmental stages of Y. tenuis. The arrangement of the elbow joint supports the view that the great appendage evolved into the chelicera of Chelicerata sensu stricto, as similar joints are found in various ingroup taxa such as Xiphosura, Opiliones or Palpigradi. With this, it also supports the interpretation of the great appendage to be homologous with the first appendage of other arthropods.     

The feeding behaviour of the Cambrian tubiculous priapulid Selkirkia

Feeding behavioural patterns have the potential to shed light on evolutionary trends, especially at the critical point of the Cambrian Explosion. In this context, we present specimens of the tubiform priapulid Selkirkia sinica, preserved in situ, collected from the lower Cambrian (Stage 3) Xiaoshiba Lagerstätte, which show the diverse morphology of its intestinal contents within the dark‐coloured gut and the formation of faecal pellets. The two digestive modes displayed by S. sinica suggest that it is a putative dietary generalist rather than a typical scavenger as previously thought. Moreover, the asymmetry exhibited by the paired caudal appendages of S. sinica provides direct evidence of the development of dorsoventral (DV) differentiation in Selkirkia, which aids our understanding of the lifestyle of this tubiculous worm.     

An early Cambrian chelicerate from the Emu Bay Shale,South Australia

The Emu Bay Shale Lagerstätte (Cambrian Series 2, Stage 4) occurs on the north coast of Kangaroo Island, South Australia. Over 50 species are known from here, including trilobites and non‐biomineralized arthropods, palaeoscolecids, a lobopodian, a polychaete, vetulicolians, nectocaridids, hyoliths, brachiopods, sponges and chancelloriids. A new chelicerate, Wisangocaris barbarahardyae gen. et sp. nov., is described herein, based on a collection of some 270 specimens. It is up to 60 mm long, with the length of the cephalic shield comprising about 30% that of the exoskeleton. The cephalic margin has three pairs of bilaterally‐symmetrical small triangular spines. A pair of small eyes is placed well forwards on the ventral margin of the cephalic shield. The trunk comprises 11 segments that increase in length while narrowing posteriorly, each possibly bearing a pair of biramous appendages; the most posterior segment is almost square whereas the others are transversely elongated. The spatulate telson is proportionately longer than in taxa such as Sanctacaris, Utahcaris and Leanchoilia. Up to eight (?four pairs) of 3 mm‐long elements bearing alternating inward‐curving short and long spines beneath the cephalic shield are interpreted as basipodal gnathobases, part of a complex feeding apparatus. A well‐developed gut includes a stomach within the cephalic shield; it extends to the base of the telson. In a few specimens there are shell fragments within the gut, including those of the trilobite Estaingia bilobata (the most common species in the biota); these fragments have sharp margins and extend across the gut lumen. The species may have been a predator or a scavenger, ingesting material already broken up by a larger predator/scavenger. The morphology of this taxon shares many overall body features with Sanctacaris, and some with Sidneyia, particularly its gnathobasic complex. These chelicerate affinities are corroborated by phylogenetic analyses.     

Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte,lower Cambrian of North Greenland

Stein, M., Peel, J.S., Siveter, D.J. & Williams, M. 2009: Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte, lower Cambrian of North Greenland. Lethaia, Vol. 43, pp. 258–265. Isoxys volucris is the most commonly occurring species in the lower Cambrian Sirius Passet Lagerstätte of North Greenland. Newly identified material allows a first, limited, account of the ventral morphology of this species, hitherto known only by the morphology of its shield. The antennula is large and robust, composed of about seven articles armed with spines, and was probably not sensorial. The postantennular limbs are serially similar, biramous with a large paddle‐shaped exopod fringed with setae. It is possible that the animal possessed a furca. The inner lamella, lining the ventral surface of the shield is recognised in Isoxys for the first time. Comparisons with other congeneric species of which aspects of the ventral morphology are known, show similarities with Isoxys auritus from China, reconsidered here, but indicate differences in antennular morphology with other species as currently understood. □Cambrian, Greenland, Isoxys, soft anatomy, Sirius Passet, palaeoecology.     

A novel respiratory architecture in the Silurian mollusc Acaenoplax

Extant aplacophorans, a group of shell‐less vermiform molluscs, respire through appendages within or projecting from a posterior cavity. Respiratory structures differ between the subclasses Caudofoveata (ctenidia within the cavity) and Solenogastres (folds of the mantle itself). Acaenoplax hayae, a Silurian vermiform mollusc from the Herefordshire Lagerstätte, England, exhibits characteristics of both these groups. While recent work places it within the crown group Aplacophora, near the caudofoveates, initial observations suggested that its respiratory structures were closer to those of the solenogastres. Here, we present new reconstructions of the posterior of Acaenoplax prepared with the aim of resolving features obscured when prior studies were undertaken. These reconstructions detail a novel posterior architecture, not closely comparable to that of either extant aplacophoran group, in which respiratory projections arise from a membrane that partly encloses a central posterior cavity. The posterior membrane is flanked by small spherical projections; both membrane and spherical projections are apparently unique within the Aplacophora. The existence of this previously undocumented respiratory system underlines the diversity of the aplacophoran clade during the Palaeozoic.