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.     

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.     

The La Voulte-sur-Rhône Lagerstätte (Middle Jurassic,France)

The La Voulte Lagerstätte (Jurassic, France) is characterized by a diverse and exceptionally well-preserved fauna. It was located along the western Tethyan margin where the submarine relief was steep. The sedimentation was dominated by marls suggesting a low energy and deep-water depositional environment. The La Voulte biota is remarkable for its biodiversity with about 60 different species. The major components are the arthropods (50% of the species), the cephalopods (10%) and the echinoderms (10%). Among the arthropods, the composition and some anatomical features suggest a relatively deep-water environment characterized by dim-light conditions. The cephalopods and other organisms (pycnogonids, asterids), have extant analogues that all live in deep-water niches always exceeding 200 m. The fauna as a whole bears the characteristic features of the present-day bathyal fauna. The La Voulte Lagerstätte may be one of the rare Jurassic Lagerstätten, if not the unique, to have fossilized a deep marine fauna.     

A 17‐element conodont apparatus from the Soom Shale Lagerstätte (Upper Ordovician), South Africa

Abstract:  Natural assemblages of a new conodont taxon, Notiodella keblon, from the Upper Ordovician Soom Shale Lagerstätte of South Africa contain 17 elements. This is the first time that a 17‐element apparatus plan has been unequivocally demonstrated in conodonts. The apparatus comprises paired P₁, P₂, P₃, M, S₁, S₂, S₃ and S₄ elements and an unpaired, axial S₀ element and provides a new template for use in the reconstruction of apparatuses from the collections of dispersed elements, particularly for those with icrion‐bearing P₁ elements and perhaps for other balognathids.     

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.     

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.     

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.     

SYNCARID CRUSTACEANS FROM THE MONTCEAU LAGERSTÄTTE (UPPER CARBONIFEROUS; FRANCE)

Abstract: Key aspects of the morphology, autecology, systematics and taphonomy of the crustacean syncarids from the Montceau Lagerstätte (Upper Carboniferous, Stephanian B; France) are presented. Palaeocaris secretanae is the most abundant faunal element of the Montceau biota and shows striking morphological similarities with Palaeocaris typus from the Mazon Creek Lagerstätte (Westphalian D; Illinois, USA). Palaeocaris secretanae was a shrimp‐like animal with a short head (no head shield), large mandibles, 14 trunk segments (the first one being reduced) and a fan‐like caudal termination. Both the body and the appendage design indicate abilities for crawling on the substratum (slender endopods) and for escape reaction (uropodal fan, pleonal flexibility), although swimming activities may have been reduced (trunk appendages with small flap‐like exopods). Details of the appendages involved in feeding, e.g. mandibles and maxillipeds, indicate poor ability for predation but point to an omnivorous detritus feeding mode. Poorly developed respiratory organs (small cylindrical epipods) suggest a relatively low level of locomotory activity. The field of vision may have been large and panoramic (stalked eyes). Rows of pores on 12 trunk segments are interpreted as possible sensory organs used for current detection. Females were brooding eggs (clusters of eggs preserved along anteroventral trunk). Microprobe analysis indicates that siderite is the major component of the nodules. Four events played a key‐role in the three‐dimensional preservation of syncarids: (1) rapid burial, (2) minimal decomposition, (3) phosphatic mineralization shortly after the animal's death and (4) nodule formation around the carcass. Palaeocaris secretanae is morphologically close to Recent syncarids such Anaspides tasmaniae (freshwater streams, Australia) in its general body plan and detailed anatomy, e.g. mouth parts, indicating morphological stasis in syncarids over more than 200 million years.     

Taphonomy of the middle Cambrian (Drumian) Ravens Throat River Lagerstätte,Rockslide Formation,Mackenzie Mountains,Northwest Territories,Canada

The Middle Cambrian (series 3, Drumian, Bolaspidella Biozone) Ravens Throat River Lagerstätte in the Rockslide Formation of the Mackenzie Mountains, northwestern Canada, contains a Burgess Shale‐type biota of similar age to the Wheeler and Marjum formations of Utah. The Rockslide Formation is a unit of deep‐water, mixed carbonate and siliciclastic facies deposited in a slope setting on the present‐day northwestern margin of Laurentia. At the fossil‐bearing locality, the unit is about 175 m thick and the lower part onlaps a fault scarp cutting lower Cambrian sandstones. It consists of a succession of shale, laminated to thin‐bedded lime mudstone, debris‐flow breccias, minor calcareous sandstone, greenish‐coloured calcareous mudstone and dolomitic siltstone, overlain by shallow‐water dolostones of the Broken Skull Formation, which indicates an overall progradational sequence. Two ~1‐m‐thick units of greenish calcareous mudstone in the upper part exhibit soft‐bodied preservation, yielding a biota dominated by bivalved arthropods and macrophytic algae, along with hyoliths and trilobites. It represents a low‐diversity in situ community. Most of the fossils occur in the lower unit, and only the more robust components are preserved. Branching burrows are present under the carapaces of some arthropods, and common millimetre‐sized disruptions of laminae are interpreted as bioturbation. The fossiliferous planar‐laminated calcareous mudstone consists of chlorite, illite, quartz silt, calcite and dolomite and is an anomalous facies in the succession. It was deposited via hemipelagic fallout of a mixture of platform‐derived and terrestrial mud. Geochemical analysis and trace‐element proxies indicate oxic bottom waters that only occasionally might have become dysoxic. Productivity in the water column was dominated by cyanobacteria. Fragments of microbial mats are common as carbonaceous seams. Complete decay of soft tissues was interrupted due to the specific sediment composition, providing support for the role of clay minerals, possibly chlorite, in the taphonomic process.     

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.     

A limulid trackway from the Late Jurassic (Tithonian) Lagerstätte of Canjuers (Var,France)

The first record of an invertebrate trackway from the lithographic limestones of the Canjuers Lagerstätte is described here. The preserved part of the trackway is approximately 39 cm long and 17 cm wide; it consists of two successive sets of imprints and the beginning of a third one. Two kinds of imprints have been recorded: the first one, well-preserved, is tetradactyl and the second one is straight, narrow and in sets of two or three imprints. The tracks described in this paper are similar to the type specimen of the ichnogenus Kouphichnium. The trackway, corresponding to locomotion behavior and left by a 26 cm wide specimen, can be related to the ichnospecies Kouphichnium lithographicum. This is the first evidence of a limulid from the Canjuers Lagerstätte.     

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.     

Multi‐jawed chaetognaths from the Chengjiang Lagerstätte (Cambrian,Series 2, Stage 3) of Yunnan,China

Chaetognaths (arrow‐worms) are enigmatic in terms of their phylogenetic position, while the existence of Protosagitta spinosa from the Chengjiang Lagerstätte suggests minimal change in their unique bodyplan since at least the early Cambrian. Apart from rare (and sometimes controversial) soft‐bodied remains, the fossil record of chaetognaths is otherwise almost entirely dependent on early Palaeozoic phosphatic microfossils, some of which are placed amongst so‐called protoconodonts. Fused spine clusters are strikingly similar to the cephalic grasping apparatus of extant forms and are assumed to have had a comparable configuration. Here we report a new chaetognath, Ankalodous sericus gen. et sp. nov., coeval with Protosagitta but with a complex feeding apparatus consisting of multiple bundles of recurved spines whose principal function appears to have been grasping. Like all other chaetognaths a predatory mode of life is likely, but its position relative to the sediment–water interface is less certain. Reduction of the feeding apparatus, from the multi‐jawed arrangement of A. sericus to the grasping spines and associated smaller teeth seen in other chaetognaths, was probably a subsequent development and conceivably was linked with a shift to a pelagic mode of life. We also report a new specimen of Protosagitta. This confirms earlier observations but it possesses hitherto unrecognized features, including a cephalic tentacle and fin rays.     

The exceptionally preserved Early Cambrian stem rhynchonelliform brachiopod Longtancunella and its implications

Zhang, Z., Holmer, L. E., Ou, Q., Han, J. & Shu, D. 2011: The exceptionally preserved Early Cambrian stem rhynchonelliform brachiopod Longtancunella and its implications. Lethaia, Vol. 44, pp. 490–495. The extraordinary Longtancunella chengjiangensis is one of the rarest and most problematic brachiopods from the Early Cambrian Chengjiang Lagerstätte, where it occurs in unique gregarious clusters with up to ten individuals attached by their stout pedicles to a single hard exoskeleton. Although the general morphology and soft anatomy of Longtancunella is available, its taxonomy and systematic placement within the Brachiopoda remains problematic. New exceptionally preserved specimens show that the pedicle of Longtancunella cannot be considered homologous with the pedicles in crown Rhynchonelliformea or Linguliformea. The pedicle of Longtancunella emerges from a ventral umbonal foramen with a rounded plate that must have been secreted by specialized ventral mantle epithelium. The rounded plate is proposed to be a homologue to the colleplax – a plate covering the umbonal foramen, which is otherwise known from the extinct rhynchonelliform chileates. Longtancunella is proposed to represent a soft‐shelled stem rhynchonelliform brachiopod with chileate features, thus demonstrating for the first time that the chileate‐like umbonal perforation functions as a pedicle opening. The unusual preservation of the annulated chileate‐like holdfast supports the view that it may have been secreted as stacks of chitinous or even mineralized ‘attachment pads’. Together with having a spirolophous lophophore and U‐shaped gut, Longtancunella displays a unique mixture of linguliform and rhynchonelliform characters that throw new light on an early stage of rhynchonelliform diversification. □Brachiopods, Cambrian, Chengjiang Lagerstätte, Chileata, colleplax, pedicle.     

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.     

Palaeoenvironment and taphonomy of a Late Jurassic (Late Tithonian) Lagerstätte from central Poland

A rich assemblage of exceptionally preserved marine and terrestrial fossils occurs in fine‐grained limestones in the upper part of the Late Tithonian (Middle Volgian) shallowing upward carbonate sequence in Central Poland. The richest horizon, a deposit known locally as the Corbulomima horizon, is named after the shallow burrowing suspension feeding bivalve Corbulomima obscura, moulds of which occur in densities of up to 500 per square metre on some bedding planes. The fauna in this bed also includes organic and phosphatic remains of a wide range of other creatures including the exuviae of limulids and decapods, disarticulated fish skeletons and rare isolated pterosaur bones and teeth. There are also perfectly preserved dragonfly wings and beetle exoskeletons. The average stable carbon and oxygen isotope values for ostracod shells and fine‐grained sediment from this horizon suggest precipitation of the calcium carbonate from warm seawater of normal marine salinity. The carbonate sediments overlying the fossiliferous horizon have been interpreted as nearshore to shoreface facies. These pass abruptly into coarse reworked intraclastic sediments interpreted as possible tsunami or storm surge over‐wash deposits. The clasts in this deposit have more positive oxygen isotope values than those in the underlying limestone, which may indicate that they were lithified in a slightly more evaporative, perhaps intertidal, setting. The succession terminates with silicified fine‐grained limestones likely to have formed in extremely shallow lagoonal environments. In contrast with the Solnhofen limestones of Lower Tithonian age in south‐central Germany the Corbulomima horizon is interpreted as a transitional deposit formed in a shallow marine setting by rapid burial with elements of both Konservat‐ and Konzentrat‐Lagerstätte preservation. □Konzentrat and Konservat‐Lagerstätte, Taphonomy, Palaeoenvironment, Paleogeography, Late Jurassic, Poland.     

A new oryctocephalid trilobite from the Balang Formation (Cambrian Stage 4) of northwestern Hunan,South China,with remarks on the classification of oryctocephalids

A new trilobite, Longaspis paiwuensis n. gen. n. sp., from the Balang Formation (Cambrian Stage 4) in northwestern Hunan, South China, is described. This rare trilobite adds to an expanding taxonomic list of organisms recognized from the Balang Lagerstätte, a deposit of exceptional preservation from the Cambrian. Longaspis paiwuensis is an unusually large-sized, micropygous oryctocephalid trilobite that has proparian facial sutures, pit-like lateral glabellar furrows, up to 17 thoracic segments, and a distinct medial notch in the pygidium; it lacks marginal spines.The classification of the family Oryctocephalidae is reviewed. Three subfamilies are recognized, and Longaspis n. gen. is assigned to the subfamily Oryctocarinae.     

Note on the gut preserved in the Lower Cambrian Lingulellotreta (Lingulata, Brachiopoda) from southern China

Lingulellotreta malongensis Rong is the earliest known taxon of the family Lingulellotretidae, which is characterized by the presence of a pedicle foramen as well as an internal pedicle tube. New material from the Early Cambrian Chengjiang Lagerstätte of southern China provides improved anatomical knowledge for lingulellotretid species especially for the digestive system. Additional gut fossils exhibit distinctly the anterior portion composed of esophagus and distended stomach, situated in the alleged visceral cavity, with the recurved intestine accommodated inside a hollow pseudointerarea. The frequency of occurrence of this intestinal layout suggests that this is not just an artefact of preservation. The gross configuration of the guts and the way they are preserved in the fossils suggest that they are in situ and, therefore, we can assume that Lingulellotreta had a hollow cavity presumably subtended by the pseuodelthyrium, which was invaded and occupied by the visceral organs. Hence, these fossils demonstrate the dangers of extrapolating crown‐group soft‐tissue configuration to the stem group.     

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.     

Chemical signatures of soft tissues distinguish between vertebrates and invertebrates from the Carboniferous Mazon Creek Lagerstätte of Illinois

The chemical composition of fossil soft tissues is a potentially powerful and yet underutilized tool for elucidating the affinity of problematic fossil organisms. In some cases, it has proven difficult to assign a problematic fossil even to the invertebrates or vertebrates (more generally chordates) based on often incompletely preserved morphology alone, and chemical composition may help to resolve such questions. Here, we use in situ Raman microspectroscopy to investigate the chemistry of a diverse array of invertebrate and vertebrate fossils from the Pennsylvanian Mazon Creek Lagerstätte of Illinois, and we generate a ChemoSpace through principal component analysis (PCA) of the in situ Raman spectra. Invertebrate soft tissues characterized by chitin (polysaccharide) fossilization products and vertebrate soft tissues characterized by protein fossilization products plot in completely separate, non‐overlapping regions of the ChemoSpace, demonstrating the utility of certain soft tissue molecular signatures as biomarkers for the original soft tissue composition of fossil organisms. The controversial problematicum Tullimonstrum, known as the Tully Monster, groups with the vertebrates, providing strong evidence of a vertebrate rather than invertebrate affinity.