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.     

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 fossils from China elucidating the phylogeny of Praesiricidae (Insecta: Hymenoptera)

A new subfamily of Praesiricidae (Pamphilioidea), Decorisiricinae subfam.n. , is erected based on three new genera: Decorisiricius gen.n. , Limbisiricius gen.n. and Brevisiricius gen.n. Two new species – Decorisiricius patulus gen. et sp.n. and D. longus sp.n. – from the Lower Cretaceous Yixian Formation and three species –Limbisiricius aequalis gen. et sp.n. , Limbisiricius complanatus sp.n. and Brevisiricius partialis gen. et sp.n. – from the Middle Jurassic Jiulongshan Formation, are described. Based on these well‐preserved new fossil specimens and previously published data, the nonmonophyly of Praesiricidae is confirmed and the phylogenetic relationships of species of Praesiricidae are analysed for the first time. Two main clades within Praesiricidae are recognized from the cladistic analysis: Decorisiricinae subfam.n. forms a monophyletic lineage, with the remaining members of Praesiricidae plus Megalodontes (Megalodontesidae) forming its sister group. The two subfamilies Archoxyelydinae and Praesiricinae are discarded with no strong supported synapomorphic characters based on phylogenetic research. A key to all genera of Praesiricidae is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:38D703ED‐127A‐4DB0‐8153‐8D78AF4AC212 .     

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.     

Two Palaeozoic Hydrothermal Vent Communities from the Southern Ural Mountains, Russia

The Sibay and Yaman Kasy massive sulphide deposits contain macrofossil assemblages that represent some of the oldest known hydrothermal vent communities. The deposits are hosted respectively by Middle Devonian and Silurian arc-related volcanic rocks in the Ural Mountains of Russia, and formed under the same environmental constraints as modern vent sulphides. The Sibay palaeocommunity comprises, in order of decreasing abundance, tubes of an indeterminate ?annelid and the vestimentiferan Tevidestus serrriformis Shpanskaya, Maslennikov and Little and articulated specimens of the modiomorphid bivalve Sibaya ivanovi gen. et sp. nov. The Yaman Kasy palaeocommunity comprises, in order of decreasing abundance, tubes of the ?polychaete Eoalvinellodes annulatus gen. et sp. nov. and the vestimentiferan Yamankasia rifeia Shpanskaya, Maslennikov and Little, and specimens of the ?kirengellid tergomyan Themoconus shadlunae gen. et sp. nov., the lingulate brachiopod Pyrodiscus lorrainae gen. et sp. nov., an indeterminate vetigastropod, and the ambonychiid bivalve Mytilarca sp. Some of these taxa have affinities to endemic taxa at modern hydrothermal vent sites and some belong to taxa that are typical of Palaeozoic non-vent marine palaeocommunities. Therefore, there has been movement of taxonomic groups in and out of the vent ecosystem through the Phanerozoic.     

Echinoderm faunas from the Lower Cretaceous (Aptian–Albian) of Alexander Island,Antarctica

Abstract: Strata assigned to the Fossil Bluff Group on Alexander Island, Antarctica, contain Aptian to Albian high‐latitude echinoderm faunas that lived at palaeolatitudes greater than 60 degrees south. The Pluto Glacier Formation, of essentially Aptian age, yields a deep‐water assemblage that includes two ophiuroids, an ophiacanthid and a representative of the ophiolepidid genus Mesophiomusium, both represented by partially articulated specimens. The echinoid fauna includes a new genus of diadematoid, Australidiadema, and a new genus of disasteroid, Notidisaster, which extend the record of both groups into the southern hemisphere. The overlying Neptune Glacier Formation, of late Albian age, yields only spatangoids which are common but rarely well enough preserved to be identified even to genus level, although at least some belong to the genus Hemiaster.     

Ophiuren (echinodermata) des hohen oberdevons im nördlichen rheinischen schiefergebirge

Four outcrops (three of which were new) in the Northern Rhenish Schiefergebirge delivered several ophiuroids from the high Famennien. Following forms are described:Protaster?ubaghsi n. sp. and a juvenile protasteroid, gen. et spec, indet., 2 new species ofFurcaster to be treated taxonomically only after revision of this genus, many specimens ofOphiaulax decheni showing plates of the primary rosette and other hitherto undescribed structures,Stephanoura belgica with many juvenile specimens, andOnychaster?velbertensis n. sp. as the first representative of the Onychasteridae in the Rhenish Schiefergebirge.     

A new,three‐dimensionally preserved enantiornithine bird (Aves: Ornithothoraces) from Gansu Province,north‐western China

In recent years, the Lower Cretaceous (Aptian) Xiagou Formation has yielded approximately 100 avian partial skeletons, many with soft‐tissue traces, from sites in the Changma Basin of Gansu Province, north‐western China. The most abundant taxon amongst these is the ornithuromorph Gansus yumenensis, but enantiornithines have also been identified in the sample. Here we describe two incomplete, semi‐articulated appendicular skeletons, the first consisting of a partial left pelvic girdle and complete pelvic limb, and the second comprised of a nearly complete right pelvic limb. Both specimens bear characteristics diagnostic of Enantiornithes, and are referred to a new taxon, Qiliania graffini gen. et sp. nov. The exceptional, three‐dimensional preservation of these specimens (compared to the crushed, nearly two‐dimensional condition of most other Early Cretaceous avian fossils) reveals new information regarding enantiornithine anatomy, evolution, and diversity. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 201–219.     

The ontogeny of cinctans (stem‐group Echinodermata) as revealed by a new genus,GraciaCystis, from the middle Cambrian of Spain

Abstract:  A new cinctan echinoderm, Graciacystis ambigua gen. et sp. nov. from Cambrian Series 3 rocks of Spain, is described based on more than 100 articulated specimens that range from 6 to 14.5 mm in thecal length. This material shows that Graciacystis ambigua, while plastic in thecal shape, is highly conservative in its thecal construction, with a fixed number of marginal plates and very limited addition of plates in the stele and ventral membrane through ontogeny. Ventral swellings on marginal elements are absent from the smallest specimens and become gradually more marked during growth. A cladistic analysis shows Graciacystis to be a basal cinctan, more derived than Sotocinctus and the Trochocystitidae and as sister group to a large clade formed by Sucocystidae + Gyrocystidae. The determinate growth pattern seen in Graciacystis seems to be the general pattern for all cinctans.     

EXCEPTIONALLY WELL-PRESERVED MIDDLE ORDOVICIAN SPONGES FROM THE LLANDEGLEY ROCKS LAGERSTÄTTE, WALES

Abstract:  A diverse fauna of three-dimensionally preserved sponges is described from nearshore volcanogenic sandstones near Llandrindod, Mid-Wales. The fauna was preserved through early marginal silicification, in rare examples with silicification of soft tissue, and includes aspicular and spicular demosponges, hexactinellids, and a heteractinid. The fauna is largely endemic, with the following new species, genera and families erected: Onerosiconcha gregalia gen. et sp. nov., Ordinisabulo quadragintaforma gen. et sp. nov., Miritubus erinaceus gen. et sp. nov., Vadosifistula milvus gen. et sp. nov., Polycornua trescelestus gen. et sp. nov. (Pseudolanciculidae fam. nov.), P. entropus sp. nov., Palaeocallyoides improbabilis gen. et sp. nov., Reticulicymbalum tres gen. et sp. nov., Triactinella rigbyi gen. et sp. nov. (Triactinellidae, fam. nov.), Spissiparies minuta gen. nov., Brevicirrus arenaceus gen. et sp. nov., Pyritonema scopula sp. nov. (Pyritonemidae, fam. nov.) and Microastraeum tenuis gen. et sp. nov. In addition, Pseudolancicula Webby and Trotter is recorded outside Australia for the first time, although only as isolated spicules. The environment represented by this fauna has not previously yielded articulated sponges, and thus the apparent endemism may be misleading.     

Comparative osteology and phylogenetic relationships of Miocaperea pulchra,the first fossil pygmy right whale genus and species (Cetacea,Mysticeti, Neobalaenidae)

A fossil pygmy right whale (Cetacea, Mysticeti, Neobalaenidae) with exquisitely preserved baleen is described for the first time in the history of cetacean palaeontology, providing a wealth of information about the evolutionary history and palaeobiogeography of Neobalaenidae. This exquisitely preserved specimen is assigned to a new genus and species, Miocaperea pulchra gen. et sp. nov. , and differs from Caperea marginata Gray, 1846, the only living taxon currently assigned to Neobalaenidae, in details of the temporal fossa and basicranium. A thorough comparative analysis of the skeleton of M. pulchra gen. et sp. nov. and C. marginata is also provided, and forms the basis of an extensive osteology‐based phylogenetic analysis, confirming the placement of M. pulchra gen. et sp. nov. within Neobalaenidae as well as the monophyly of Neobalaenidae and Balaenidae; the phylogenetic results support the validity of the superfamily Balaenoidea. No relationship with Balaenopteroidea was found by the present study, and thus the balaenopterid‐like morphological features observed in C. marginata must have resulted from parallel evolution. The presence of M. pulchra gen. et sp. nov. around 2000 km north from the northernmost sightings of C. marginata suggests that different ecological conditions were able to support pygmy right whale populations in what is now Peru, and that subsequent environmental change caused a southern shift in the distribution of the living neobalaenid whales. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 876–911.     

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.     

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.     

A new brittle star (Ophiuroidea: Ophiodermatina) from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

Knowledge on the morphology of Early Triassic ophiuroids is very limited, in spite of the relatively high number of described species. This hampers attempts to use morphology-based phylogenetic analyses in order to explore the early diversification of the ophiuroid modern clades between the latest Paleozoic and the early Mesozoic. Here, a new ophiuroid from the Early Triassic of southern Idaho (USA) is described and assigned to the new taxon Shoshonura brayardi Thuy nov. gen., nov. sp. It is part of an exceptional Early Triassic faunal assemblage recently described as the Paris Biota. On account of the arm morphology and the shape of the arm spines and arm spine articulations, Shoshonura brayardi nov. gen., nov. sp. is considered a basal member of the suborder Ophiodermatina.     

Problematic megafossils in Cambrian palaeosols of South Australia

Abstract: Red calcareous Middle Cambrian palaeosols from the upper Moodlatana Formation in the eastern Flinders Ranges of South Australia formed in well‐drained subhumid floodplains and include a variety of problematic fossils. The fossils are preserved like trace fossil endichnia but do not appear to be traces of burrows or other animal movement. They are here regarded as remains of sessile organisms, comparable with fungi or plants living in place, and are formally named as palaeobotanical form genera under provisions of the International Code of Botanical Nomenclature. Most common are slender (0.5–2 mm) branching filaments flanked by green‐grey reduction haloes within the red matrix of palaeosol surface horizons (Prasinema gracile gen. et sp. nov.). Other axial structures (Prasinema nodosum and P. adunatum gen. et spp. nov.) are larger and show distinctive surface irregularities (short protuberances and irregular striations, respectively). The size and form of these filaments are most like rhizines of soil‐crust lichens. Other evidence of life on land includes quilted spheroids (Erytholus globosus gen. et sp. nov.) and thallose impressions (Farghera sp. indet.), which may have been slime moulds and lichens, respectively. These distinctive fossils in Cambrian palaeosols represent communities comparable with modern biological soil crusts.     

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).     

Mass mortality of an asteriid starfish (Forcipulatida,Asteroidea, Echinodermata) from the late Maastrichtian (Late Cretaceous) of Morocco

Abstract: A mass occurrence, numbering tens of thousands of individuals in a single lens, belonging to a single species of asteroid, is described from the late Maastrichtian (Late Cetaceous) of Morocco. The lens of partially silicified asteroidal limestone is made up largely of fully articulated specimens of similar size and probably represents the mass mortality of a single recruitment. By comparison with mass strandings of the present‐day species Asterias rubens (Linnaeus), it can be inferred that a feeding swarm of individuals was swept into a submarine channel by either a storm or an exceptionally strong tidal current, and permanently buried. The genus and species are herein described as Cretasterias reticulatus gen. et sp. nov. The exceptional preservation of the material enables the identification of wreath organs (clusters of crossed pedicellariae set in a dermal pad around spines) for the first time in the fossil record. Comparison between extant Asteriidae, putative fossil asteriids and C. reticulatus provides ambiguous evidence of its affinities; it appears to display a combination of plesiomorphic and derived characters. It is shown that all Mesozoic forcipulatid asteroids described so far share a very simple arm construction (single row of adradial ossicles) unknown in adult extant Forcipulatida.     

LATE JURASSIC PALAEONTINIDS (HOMOPTERA: AUCHENOR-RHYNCHA) FROM HEBEI AND LIAONING PROVINCES IN CHINA*

Abstract Two new genera and six new species of Palaeontinidae are named and described: Liaocossus hui gen. et sp. nov., L. beipiaoensis gen. et sp. nov., L. exiguus gen. et sp. nov., L. fengningensis gen. et sp. nov., L. pingquanensis gen. et sp. nov., and Miracossus ingentius gen. et sp. nov., The entire body of these new fossils are perfectly preserved and enable us to fix accurately the body characteristics of primitive palaeontinids. All the specimens were collected from Late Jurassic Yixian Formation of Northern Hebei and Western Liaoning Provinces in China.