A cryptic record of Burgess Shale‐type diversity from the early Cambrian of Baltica

Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).     

Small carbonaceous fossils (SCFs) from the Terreneuvian (lower Cambrian) of Baltica

We describe a new assemblage of small carbonaceous fossils (SCFs) from diagenetically minimally altered clays and siltstones of Terreneuvian age from the Lontova and Voosi formations of Estonia, Lithuania and Russia. This is the first detailed account of an SCF assemblage from the Terreneuvian and includes a number of previously undocumented Cambrian organisms. Recognizably bilaterian‐derived SCFs include abundant protoconodonts (total‐group Chaetognatha), and distinctive cuticular spines of scalidophoran worms. Alongside these metazoan remains are a range of protistan‐grade fossils, including Retiranus balticus gen. et sp. nov., a distinctive funnel‐shaped or sheet‐like problematicum characterized by terminal or marginal vesicles, and Lontohystrichosphaera grandis gen. et sp. nov., a large (100–550 μm) ornamented vesicular microfossil. Together these data offer a fundamentally enriched view of Terreneuvian life in the epicratonic seas of Baltica, from an episode where records of non‐biomineralized life are currently sparse. Even so, the recovered assemblages contain a lower diversity of metazoans than SCF biotas from younger (Stage 4) Baltic successions that represent broadly equivalent environments, echoing the diversification signal recorded in the coeval shelly and trace‐fossil records. Close comparison to the biostratigraphical signal from Fortunian small shelly fossils supports a late Fortunian age for most of the Lontova/Voosi succession, rather than a younger (wholly Stage 2) range.     

Diverse middle Ordovician palaeoscolecidan worms from the Builth‐Llandrindod Inlier of central Wales

Abstract: Palaeoscolecidan worms are rare, Early Palaeozoic fossils with uncertain affinities within the Ecdysozoa. They are locally abundant in the Cambrian and scattered in the Early Ordovician, but very sparse thereafter. Forty‐four specimens have been collected from the Middle Ordovician of the Builth‐Llandrindod Inlier of Mid Wales and include well‐preserved material assigned to seven new genera, with four additional species in open nomenclature. An additional specimen from the Arenig Pontyfenni Formation of South Wales is also described in open nomenclature. The total demonstrates much greater palaeoscolecid diversity than hitherto suspected for this time. The specimens are preserved as cuticle fragments in shales and siltstones, often of submillimetre size but in many cases with excellent preservation. The level of detail preserved in some is equal to that found in Cambrian phosphatized faunas. The new approach to collecting, and the recognition that this material can yield taxonomically useful information, opens new avenues for palaeoscolecidan research in siliciclastic environments. The new taxa are the following: Radnorscolex bwlchi gen. et sp. nov., Aggerscolex murchisoni gen. et sp. nov., Bullascolex inserere gen. et sp. nov., Wernia eximia gen. et sp. nov., Ulexiscolex ormrodi gen. et sp. nov., Pluoscolex linearis gen. et sp. nov. and Loriciscolex cuspidus gen. et sp. nov. The high diversity, and the taxonomic separation from known groups described primarily from Cambrian carbonates, implies that palaeoscolecidans either diversified significantly during the Ordovician or were taxonomically segregated between carbonate and siliciclastic settings. Palaeobiological findings also include confirmation that some palaeoscolecid basal cuticles were solid and others reticulate, plates (and platelets) could form by lateral accretion, plates were in part primarily phosphatic and in part organic and that in at least some groups, platelet secretion occurred external to plate secretion.     

PERMIAN OSTRACODS FROM THE LERCARA FORMATION (MIDDLE TRIASSIC–CARNIAN?), SICILY,ITALY

Abstract: A rich, diverse Permian ostracod fauna has been recovered from the red and grey, laminated shales and quartz‐rich shales of the Triassic Lercara Formation. Forty‐seven species have been identified, 13 of which are newly described here; they belong to 26 genera of which three are new: Anahuacia lercaraensis sp. nov., Aurigerites siciliaensis sp. nov., Bairdia portellaensis sp. nov., Cristanaria? katyae sp. nov., Fabalicypris gruendeli sp. nov., Lethiersa salomonensis gen. et sp. nov., Lethiersia sinusoventralis gen. et sp. nov., Portella trapezoida gen. et sp. nov., Siciliella elongata gen. et sp. nov., Siciliella infernespinosa gen. et sp. nov., Siciliella prima gen. et sp. nov., Siciliella quadrata gen. et sp. nov., and Siciliella spinorobusta gen. et sp. nov. The assemblages contain or are composed of palaeopsychrospheric forms, which are regarded as index fossils for deep environments. The bathymetry of the different associations in life is evaluated.     

New data on molluscs and their shell microstructures from the Middle Cambrian Gowers Formation, Australia

Abstract: Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation (Ptychagnostus atavus/Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi‐nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite‐shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre‐like microstructures in many of these fossils reveals how widespread these strong (fracture‐resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov.     

Palaeobiology of tanaidaceans (Crustacea: Peracarida) from Cretaceous ambers: extending the scarce fossil record of a diverse peracarid group

Diverse assemblages of tanaidacean peracarid crustaceans from western Tethyan continental deposits suggest that the group was relatively common in or around ancient resin‐producing forests. Here we report the results of an examination of 13 tanaidacean specimens from three Cretaceous (Albian to Turonian) French amber deposits. Two new species of the fossil family Alavatanaidae are placed in the previously described Early Cretaceous genus Eurotanais: Eurotanais pyrenaensis sp. nov. from Cenomanian Pyrenean amber (Fourtou, Aude) and Eurotanais seilacheri sp. nov. from Turonian Vendean amber (La Garnache, Vendée). The remaining specimens are placed in three newly erected genera and species (but family incertae sedis): Arcantitanais turpis gen. et sp. nov. from Albian–Cenomanian Charentese amber (Archingeay, Charente‐Maritime), and Tytthotanais tenvis gen. et sp. nov. and Armadillopsis rara gen. et sp. nov. from Pyrenean amber. These are the first formally described fossils that might be related to the paratanaoidean families Nototanaidae and Paratanaidae, sharing with these some putatively derived features and providing possible evidence for the antiquity and morphological stability of these families and the suborder Tanaidomorpha. The distinctive features and character combinations of these fossil taxa are discussed in connection with possible relationships to the living lineages of tanaidaceans. Propagation phase‐contrast X‐ray synchrotron microtomography was used to obtain high‐quality 3D images for some fossils. A discussion is provided on the putative palaeobiology of tanaidaceans and the French resiniferous forest ecosystem. The discovery of these new tanaidaceans extends the palaeogeographical distribution and stratigraphical range of the family Alavatanaidae and sheds new light on the palaeoecology and diversity of tanaidaceans in pre‐angiospermous woodlands.     

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.     

New palaeoscolecid worms from the Furongian (upper Cambrian) of Hunan,South China: is Markuelia an embryonic palaeoscolecid?

Abstract: Three‐dimensional fragments of palaeoscolecid cuticle have been recovered from the Furongian (upper Cambrian) of Hunan, South China. Extraordinary preservation of the fossils shows exquisite surface details indicating a three‐layered structure of the cuticle. One new genus and two new species Dispinoscolex decorus gen. et sp. nov. and Schistoscolex hunanensis sp. nov. are described. The co‐occurrence of these palaeoscolecid remains with those of Markuelia hunanensis allowed us to test the hypothesis that Markuelia, known hitherto only from embryonic remains, is an embryonic palaeoscolecid. The comparative anatomy of Markuelia and the co‐occurring palaeoscolecids shows a number of distinctions, particularly in the structure of the tail; all similarities are scalidophoran or introvertan (cycloneuralian) symplesiomorphies. The available evidence does not support the interpretation of Markuelia as an embryonic palaeoscolecid.     

A new family of Cambrian rhynchonelliformean brachiopods (Order Naukatida) with an aberrant coral‐like morphology

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian (Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian (Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral‐like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube‐like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian (Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef‐building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well‐developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle.     

The oldest cinctan carpoid (stem‐group Echinodermata), and the evolution of the water vascular system

A new cinctan ( Protocinctus mansillaensis gen. et sp. nov. ), from the early Middle Cambrian of the Iberian Chains (north‐east Spain), is described with the aid of X‐ray microtomography and three‐dimensional computer models. Investigation in this manner was possible because of the unusual condition of the fossils, which are preserved as recrystallized calcite. Protocinctus gen nov. possesses an elongate body with a single left anterior feeding groove and an open posterior marginal frame (in ventral view): this combination of characters is unique amongst cinctans. Through the study of original specimens and ‘virtual fossils’ it was possible to reconstruct the palaeobiology of Protocinctus gen. nov. : cinctans are interpreted as pharyngeal basket feeders with a U‐shaped gut, using their posterior appendage to aid stability on the sediment surface. Cinctans are critical to understanding the evolutionary history of the echinoderm stem group, as they illustrate the transition from a paired water vascular system (basal) to one constructed from just the left hydrocoel (derived). The former condition is also observed in another group of stem‐group echinoderms, the ctenocystoids, and is hypothesized for the latest common ancestor of the echinoderms and hemichordates.     

Records of bizarre Jurassic brachycerans in the Daohugou biota,China (Diptera,Brachycera, Archisargidae and Rhagionemestriidae)

Abstract: Four new Brachycera fossils were collected from the Daohugou biota, China. Among these, two impressions demonstrating peculiar wing venation can be designated as two new species of a new genus (Mostovskisargus portentosus gen. et sp. nov. and M. signatus sp. nov.) referred to a new subfamily Mostovskisarginae (subfam. nov.) within Archisargidae. The third impression belongs to a new species (Calosargus (Pterosargus) sinicus sp. nov.) referred to the subgenus Pterosargus Mostovski, 1997 of Calosargus Mostovski, 1997 within Archisarginae, Archisargidae. It is only the second member of the subgenus Pterosargus worldwide. The fourth impression is a nearly complete fly, Jurassinemestrinus orientalis gen. et sp. nov. referable to Rhagionemestriidae. It reveals new morphological data about rhagionemestriids and extends the geographical distribution of this group beyond Europe and Central Asia during the Mesozoic. A correlation for the archisargid composition of taxa in Daohugou and Karabastau formations is discussed. The age of the fly‐bearing strata is briefly reassessed and can be more accurately limited to the Callovian–Oxfordian based on both biostratigraphical correlation and radiometric dating.     

New Genera and Species of Psychodoid Flies from the Lower Cretaceous Amber Lebanon

Until now, only two Psychodoidea were known from Lebanese amber. We describe two new genera and species of Phlebotomidae ( Mesophlebotomites hennigi gen. et sp. nov., Libanophlebotomus lutfallahi gen. et sp. nov.) and four new genera with six new species of Psychodidae ( Paleopsychoda solignaci gen. et sp. nov., Paleopsychoda jacquelinae sp. nov., Protopsychoda nadiae gen. et sp. nov., Protopsychoda hammanaensis sp. nov., Libanopsychoda abillamai gen. et sp. nov., Cretapsychoda inexpectata gen. et sp. nov.) from the Lower Cretaceous amber of Hammana/Mdeirij, Lebanon. These fossils are included in a phylogenetic analysis of the subfamilies of Psychodoidea. This superfamily was probably as diverse in the Early Cretaceous as now.     

Chondrichthyan‐like scales from the Middle Ordovician of Australia

Abstract: Microvertebrate sampling of the Stairway Sandstone (Darriwilian, Middle Ordovician, central Australia) has yielded scales that are chondrichthyan‐like in their overall construction, and Tantalepis gatehousei gen. et sp. nov. is erected here to describe these specimens. Tantalepis gatehousei gen. et sp. nov. is the stratigraphically oldest microsquamous taxon described thus far, and the ‘shark‐like’ appearance of the scales may extend the chondrichthyan lineage back into the Middle Ordovician. The presence of ‘shark‐like’ scales in the fossil record some 50 myr prior to the first articulated chondrichthyan body fossils and 44 myr before the first clearly identifiable chondrichthyan teeth suggests there is a considerable scope for the recovery of articulated specimens with which to document the early history of crown gnathostomes. Traditional hypotheses of phylogenetic relationships among early jawed vertebrates were recently challenged by the proposal of a radically different tree topology. However, the development of a new data set specifically addressing scale‐based characters is required before taxa such as Tantalepis, that are based upon disarticulated remains alone, can be firmly placed within the emerging, revised, evolutionary narrative.     

New fossil mantidflies (Insecta: Neuroptera: Mantispidae) from the Mesozoic of north‐eastern China

Abstract:  Three new genera and four new species of the extinct mantidfly subfamily Mesomantispinae (Insecta: Neuroptera: Mantispidae) are described from the Lower Cretaceous Yixian Formation of Liaoning and the Middle Jurassic Jiulongshan Formation of Inner Mongolia: Archaeodrepanicus nuddsi gen. et sp. nov., A. acutus gen. et sp. nov., Sinomesomantispa microdentata gen. et sp. nov., (Yixian Formation) and Clavifemora rotundata gen. et sp. nov. (Jiulongshan Formation). The specimens described herein represent the first Mesozoic mantidfly compression fossils to have body parts preserved, including the specialized raptorial forelegs articulated to the prothorax anteriorly, an autapomorphy of the family. These new taxa further confirm the placement of the subfamily Mesomantispinae within the family Mantispidae; however, the monophyly of Mesomantispinae has not been confirmed, and it is likely that it will prove to be paraphyletic.     

New archaeocyatha from the Atdabanian Stage of the Siberian Platform

New Archaeocyatha genera and species, Turgorocyathus elegans gen. et sp. nov., Angustocyathus porus gen. et sp. nov., Flossocyathus squamosus gen. et sp. nov., Rotundocyathus oimuranicus sp. nov., and Carinacyathus inopinatus sp. nov., from the type sections of the Atdabanian Stage of the Lower Cambrian of the Siberian Platform are described.     

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.     

Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte,North Greenland,and the polarity of morphological characters in basal deuterostomes

Abstract: Vetulicolians are problematic Cambrian fossils with a debated phylogenetic history. Here, we describe two vetulicolian specimens from the Lower Cambrian Sirius Passet locality in North Greenland. One of the specimens is assigned to Ooedigera peeli gen. et sp. nov, whereas the other is retained under open nomenclature. The mode of tail flexibility has been debated in the literature, and we argue here that the tail normally flexed laterally to generate power strokes rather than dorsoventrally. The phylogenetic relationships of vetulicolians are discussed in the light of current knowledge of deuterostome phylogeny and morphology, and it is concluded that the best hypothesis on currently available evidence is that vetulicolians are a clade or paraphyletic assemblage of stem‐Deuterostomata. The presence of a voluminous, sediment‐filled anterior chamber suggests that the atrium may be a synapomorphy of deuterostomes.     

A palaeoscolecid worm from the Burgess Shale

Palaeoscolecid worms are a ubiquitous group of Early Palaeozoic ecdysozoans that are curiously lacking in the archetypal Cambrian Lagerstätten, the Burgess Shale. Here I describe Scathascolex minor gen. et sp. nov, the first unequivocal palaeoscolecid from this site. Scathascolex is armoured with simple Hadimopanella‐like plates, but lacks smaller platelets, pointing to a close affinity with the Palaeoscolecida sensu stricto. Neither preservational nor environmental factors account for the scarcity of palaeoscolecids in the Burgess Shale, which presumably represents an ecological phenomenon.