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.     

Experimental investigation of insect deposition in lentic environments and implications for formation of Konservat Lagerstätten

Terrestrial insects are often remarkably well preserved in lacustrine Konservat Lagerstätten. However, the assumption that carcasses should sink fast through the water column seems contradictory as this scenario is unlikely due to excessive buoyancy and surface tension. The mechanisms that promote rapid and permanent emplacement onto the sediment surface (RPESS) of such terrestrial animal remains are not fully understood. Here we use taphonomic experiments to show that floating in water, growth of microbial biofilms and reception of rapid sediment load promote RPESS of terrestrial insect remains in lentic water bodies. Our results show that the optimum conditions for RPESS occur when terrestrial insects enter a lentic water body in articulation, experience brief decay in association with growth of microbes, then are buried rapidly by airborne volcanic ash. These results provide a model for preservation of articulated terrestrial insects and emphasize the importance of microbial activity and volcanism for insect preservation in lacustrine Konservat Lagerstätten.     

Unveiling biases in soft‐tissue phosphatization: extensive preservation of musculature in the Cretaceous (Cenomanian) polychaete Rollinschaeta myoplena (Annelida: Amphinomidae)

The process of soft‐tissue phosphatization (the replication of labile tissues by calcium phosphate) is responsible for many instances of high‐resolution soft tissue preservation, often revealing anatomical insights into the animals that so preserved. However, while much work has gone into exploring key issues such as biases and micro‐controls, phosphatization remains poorly understood as a taphonomic process. Here, using camera lucida, plain‐light microscopy and SEM imagery, we address this issue by describing the taphonomy and fidelity of the musculature of Rollinschaeta myoplena Parry et al., a phosphatized annelid from the Cretaceous Konservat‐Lagerstätten of Hakel and Hjoula, Lebanon, with an unprecedented quantity of three‐dimensional soft‐tissue preservation. Analysis highlights two strong, previously recognized biases affecting the process of phosphatization: (1) a taxonomic bias restricted to R. myoplena that triggers unusually extensive phosphatization; and (2) a tissue bias whereby longitudinal and parapodial musculature show markedly higher fidelity in comparison to the musculature of the intestine and body wall circular muscles. Potential explanations for these biases include internal phosphate‐enrichment by relative muscle density, the relative rate of decay and the physiology of musculature. Incongruence between experimental decay series for polychaetes and the prevalence of labile tissue preservation over recalcitrant tissues in R. myoplena exposes the limits of decay experiments for understanding exceptional preservation.     

Fossil representatives of the family Greenideidae (Hemiptera, Aphidoidea) from the Miocene of Europe

Four new aphid species (Hemiptera, Aphidoidea, Greenideidae) are described on the basis of imprints in oil-shales from the Miocene of Europe (Rubielos de Mora, Spain; Vishnevaya Balka ?Stavropol?, Russia): Eutrichosiphum europaeum nov. sp., Greenidea hispanica nov. sp., Greenideoida (Pentatrichosiphum) turolensis nov. sp. and Mollitrichosiphum rubusensis nov. sp. All the taxa are placed within recent genera of the subfamily Greenideinae. Miocene representatives of this subfamily have been found in the south of Europe. Later changes of climatic conditions limited its geographic range to south-eastern Asia, where it is now represented by over 130 species of seven genera.     

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.     

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.     

Exceptionally preserved tadpoles from the Miocene of Libros,Spain: ecomorphological reconstruction and the impact of ontogeny upon taphonomy

McNamara, M.E., Orr, P.J., Kearns, S.L., Alcalá, L., Anadón, P. & Peñalver‐Mollá, E. 2010: Exceptionally preserved tadpoles from the Miocene of Libros, Spain: ecomorphological reconstruction and the impact of ontogeny upon taphonomy. Lethaia, Vol. 43, pp. 290–306. The Libros exceptional biota from the Upper Miocene of NE Spain includes abundant frog tadpoles (Rana pueyoi) preserved in finely laminated lacustrine mudstones. The tadpoles exhibit a depressed body, short tail, low tail fins, dorso‐laterally directed eyes and jaw sheaths; these features identify the Libros tadpoles as members of the benthic lentic ecomorphological guild. This, the first ecomorphological reconstruction of a fossil tadpole, supports phylogenetic evidence that this ecology is a conserved ranid feature. The soft‐tissue features of the Libros tadpoles are characterized by several modes of preservation. The space occupied previously by the brain is defined by calcium carbonate, the nerve cord is defined by calcium phosphate, and jaw sheaths and bone marrow are preserved as organic remains. Gut contents (and coprolites adjacent to specimens) comprise ingested fine‐grained sedimentary detritus and epiphyton. The body outline and the eyespots, nares, abdominal cavity, notochord, caudal myotomes and fins are defined by a carbonaceous bacterial biofilm. A similar biofilm in adult specimens of R. pueyoi from Libros defines only the body outline, not any internal anatomical features. In the adult frogs, but not in the tadpoles, calcium phosphate and calcium sulphate precipitated in association with integumentary tissues. These differences in the mode of preservation between the adult frogs and tadpoles reflect ontogenetic factors. □Anuran, ecology, soft‐tissue, tadpoles, taphonomy.     

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.     

Early Miocene repetitive vegetation and climatic changes in the lacustrine deposits of the Rubielos de Mora Basin (Teruel, NE Spain)

Pollen analysis of a Lower Miocene succession from the Rubielos de Mora Basin (NE Spain) has been carried out with the aim of reconstructing the flora, vegetation and climatic changes. Previous paleobotanical studies on these sedimentary rocks and adjacent areas interpreted very diverse climates for the Early Miocene: from humid temperate to dry subtropical. In this study, a rich thermophilous pollen spectrum and a diverse subarid flora including Nitraria, Caesalpiniaceae, Ephedra and Acacia, indicative of a dry subtropical climate are identified. On the other hand, mesothermic taxa with high water requirements are also abundant. Therefore, the pollen assemblages evidence the juxtaposition of very contrasted environments: the presence of subdesertic taxa, typical of plants growing in the lowlands and conditioned by a long warm, dry season, together with others with very high water requirements, needing constant water. This can be explained by the presence of the Rubielos de Mora Lake providing local conditions for developing riparian forests. The vegetation was clearly controlled by the water availability under a subtropical and dry-seasonal climate. Pollen changes along the succession, which coincide with sedimentological changes, are related to climatic variations. Alternation in pollen taxa (thermophilous–dry vs. mesothermic–riparian) reflects the influence of the cyclicity of temperature and precipitation on the lake level and vegetation.     

The Upper Carboniferous Hamilton Fossil-Lagerstätte in Kansas: a valley-fill, tidally influenced deposit

A diverse assemblage of unusually well-preserved marine, euryhaline, freshwater, and terrestrial fossils (invertebrates, vertebrates, and plants) occurs within an Upper Carboniferous (Stephanian) Konservat Fossil-Lagerstätte near Hamilton, Kansas, USA. The Lagerstätte occurs within a paleovalley that was incised into the surrounding Carboniferous cyclothemic sequence during a time of low sea level, and was then filled-in during a subsequent transgression. The stratigraphically lowest and most voluminous facies within the valley is a cross-bedded, polymictic limestone conglomerate that contains caliche clasts and charcoal fragments as well as some normal-marine fossils apparently in situ. The origin of the conglomerate is enigmatic, but it was probably deposited by a migrating tidal channel. Overlying and interbedded with the conglomerate is an ostracode wackestone that contains plants (primarily seed ferns and ferns), eurypterids, shrimp, brachiopods, bivalves, and rare fish. The ostracode wackestone was deposited in a low-energy, marginal-marine environment. A thin sequence (<1 m thick) of interbedded laminated limestone and mudstone overlies the conglomerate in a small area. This facies contains a well-preserved mixed assemblage of terrestrial (conifers, insects, myriapods, reptile), freshwater (ostracodes), aquatic (amphibians, reptile), brackish or euryhaline (ostracodes, eurypterids, spirorbids, fish), and marine (brachiopods, echinoderms) fossils. Many of the vertebrates are articulated and show no evidence of preburial decay, scavenging, or predation. A few vertebrates exhibit signs of flotation. Most articulated vertebrate specimens exhibit soft-tissue preservation in the form of dark-brown to black early-diagenetic microbialite body outlines (‘skin preservation’) containing fossil bacteria. Rhythmic patterns of lamination thickness variation in the limestones and mudstones indicate that this facies was deposited in a tidal environment. High sedimentation rate and variable salinity (and therefore exclusion of bioturbators and invertebrate scavengers) are interpreted as key elements that led to the excellent preservation of the fossils in this ancient estuarine environment. □Lagerstätte, taphonomy, estuarine, tidal bedding, paleovalley, Carboniferous, Kansas.     

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.     

Skeletal completeness of the non‐avian theropod dinosaur fossil record

Non‐avian theropods were a highly successful clade of bipedal, predominantly carnivorous, dinosaurs. Their diversity and macroevolutionary patterns have been the subject of many studies. Changes in fossil specimen completeness through time and space can bias our understanding of macroevolution. Here, we quantify the completeness of 455 non‐avian theropod species using the skeletal completeness metric (SCM), which calculates the proportion of a complete skeleton preserved for a specimen. Temporal patterns of theropod skeletal completeness show peaks in the Carnian, Oxfordian–Kimmeridgian and Barremian–Aptian, and lows in the Berriasian and Hauterivian. Lagerstätten primarily drive the peaks in completeness and observed taxonomic diversity in the Oxfordian–Kimmeridgian and the Barremian–Aptian. Theropods have a significantly lower distribution of completeness scores than contemporary sauropodomorph dinosaurs but change in completeness through time for the two groups shows a significant correlation when conservation Lagerstätten are excluded, possibly indicating that both records are primarily driven by geology and sampling availability. Our results reveal relatively weak temporal sampling biases acting on the theropod record but relatively strong spatial and environmental biases. Asia has a significantly more complete record than any other continent, the mid northern latitudes have the highest abundance of finds, and most complete theropod skeletons come from lacustrine and aeolian environments. We suggest that these patterns result from historical research focus, modern climate dynamics, and depositional transportation energy plus association with conservation Lagerstätten, respectively. Furthermore, we find possible ecological biases acting on different theropod subgroups, but body size does not influence theropod completeness on a global scale.     

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.     

The Montceau-les-Mines Lagerstätte (Late Carboniferous,France)

The Montceau-les-Mines Lagerstätte (Late Stephanian, Late Carboniferous) is located northeast of the French Massif Central. Situated at equatorial latitudes during the Pennsylvanian, this Lagerstätte, probably a freshwater environment, preserves a rich and diverse flora (lycopsids, sphenopsids, ferns, pteridosperms, and cordaites) and fauna (bivalves, annelids, crustaceans, myriapods, insects, chelicerates, myxinoids, actinopterygians, sarcopterygians and tetrapods). These exceptionally preserved fossils can be found either flattened in shales or three-dimensionally preserved in sideritic nodules. The fossils from the nodules are exceptional for at least two reasons: the absence of major disarticulation of their body structure and the preservation of soft parts and extremely fragile cuticular structures. Such preservation was made possible by the combination of several factors: rapid burial in fine anoxic mud, early siderite precipitation (inducing the nodule formation) and phosphatization of cuticles and soft-bodied features.     

Buoyancy mechanisms limit preservation of coleoid cephalopod soft tissues in Mesozoic Lagerstätten

Coleoid cephalopods are characterized by internalization of their shell, and are divided into the ten‐armed Decabrachia (squids and cuttlefish) and the eight‐armed Vampyropoda (octopuses and vampire squid). They have a rich fossil record predominantly of the limited biomineralized skeletal elements they possess: arm hooks, statoliths, mouthparts (the buccal mass) and internal shell (gladius or pen), although exquisitely preserved soft tissue coleoids are known from several Lagerstätten worldwide. Recent studies have shown that although morphological similarities between extant decabrachian gladii and fossil examples exist, no known examples of fossil decabrachians are currently known. However, molecular clock data and phylogenetic bracketing suggest that they should be present in Lagerstätten that are rich in vampyropod soft tissue fossils (i.e. Hâkel and Hâdjoula Lagerstätten, Cretaceous, Lebanon). We propose that a hitherto unknown taphonomic bias pertaining to the differing methods of buoyancy control within coleoid groups limits preservation potential. Both negatively and neutrally buoyant decabrachians use chemical buoyancy control (ammonia) whereas vampyropods do not. In the event of rapid burial in an environment conducive to exceptional preservation, ammonia dramatically decreases the ability of the decabrachian carcass to generate the required pH for authigenic calcium phosphate replacement, limiting its preservation potential. Moreover, the greater surface area and comparatively fragile dermis further decrease the potential for fossilization. This taphonomic bias may have contributed to the lack of preserved labile soft‐tissues in other cephalopods groups such as the ammonoids.     

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.     

High‐resolution δ13Corg chemostratigraphy links the Decorah impact structure and Winneshiek Konservat‐Lagerstätte to the Darriwilian (Middle Ordovician) global peak influx of meteorites

The precise age of the Winneshiek Shale, a recently discovered Konservat‐Lagerstätte located in a very unusual depositional setting inside the Decorah impact structure, has remained uncertain in the absence of biostratigraphically highly diagnostic fossils. This chemostratigraphical study, based on δ¹³C_org data from 36 drill core samples through the shale, shows that the age ranges from the upper part of a small unnamed δ¹³C excursion in the Dw1 Stage Slice of the Darriwilian Global Stage to the lower part of the MDICE excursion in Stage Slice Dw2 of the same stage. This Dw1–Dw2 interval has an isotopic age of ~464–467 Ma. The gradational contact between the Winneshiek Shale and the underlying, rapidly deposited, impact breccia indicates minimal time difference between the impact event and the Winneshiek Shale. New age data show that the Decorah impact event was coeval with the early Darriwilian abnormally high influx of micrometeorites and meteorites recorded in sections in Baltoscandia, Russia and China and that the Decorah crater can be included among the unusually large number of meteorite craters formed during Middle and early Late Ordovician time. As is commonly the case in black shale deposits, the partly uniquely preserved Winneshiek Shale crater fauna is impoverished taxonomically and adds numerically relatively little to the conspicuous and much discussed Darriwilian global biodiversification increase.     

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.