QUALITY OF THE TRIASSIC–JURASSIC BIVALVE FOSSIL RECORD IN NORTHWEST EUROPE

Abstract: The quality of the Triassic–Jurassic bivalve fossil record in northwest Europe has been measured using the Simple Completeness Metric (SCM). The SCM has been applied to the fossil record of total bivalve diversity and to the records of different ecological guilds. The Westbury and Lilstock Formations record high SCM values for most ecological groups. The ‘Pre‐Planorbis Beds’ of the lower Lias Group, however, witness a precipitous decline in the completeness of most guilds and emigration of taxa due to localized marine anoxia is a likely cause. Neither variation in lithofacies, shell mineralogy, sedimentary rock outcrop area, nor sequence architecture can convincingly explain the observed patterns of completeness. Our SCM data reveal that the Early Jurassic fossil record of infaunal suspension‐feeding bivalves is significantly poorer than that of epifaunal bivalves. Any differences in the apparent Rhaetian extinction rates between these two guilds should therefore be viewed with caution. Analyses of selectivity during the Late Triassic mass extinction based on studies of global databases appear robust in light of our SCM data. Nevertheless, future investigations of the Triassic–Jurassic benthic marine ecosystem undertaken at a finer‐resolution, may need to account for the poor quality of the Early Jurassic fossil records of certain ecological guilds, such as the infaunal suspension‐feeding taxa.     

Trace fossil associations related to facies of an upper Ordovician low wave energy shoreface and shelf, Oslo-Asker district, Norway

Three trace fossil associations have been identified from facies interpreted as low wave energy upper, middle and lower shoreface deposits, transitional shoreface/shelf deposits and open epicontinental shelf deposits. These were developed in pre-regressive, regressive and transgressive phases associated with eustatic sea level changes caused by the upper Ordovician glaciation. The three associations are the Skolithos-Diplocraterion association, the Diplichnites-Phycodes association and the Thalassinoides association. The Skolithos-Diplocraterion association inhabited upper, middle and lower shoreface environments, the Diplichnites-Phycodes association a lowermost shoreface environment and the Thalassinoides association inhabited offshore epicontinental shelf environmcnts. The Skolithos-Diplocraferion and Diplichnites-Phycodes associations bear strong affinities with previously described Skolithos ichnofacies assemblages (indicating shoreline environments) and the Cruziana ichnofacies forms (indicating shallow water environments down to wave base) respectively. The Thalassinoides assemblage, however, closely compares with assemblages described from Mesozoic epicontinental shelf sequences. Trace fossil diversity achieves a maximum in the lowermost shoreface environment and reduces in deeper water and shallower water environments. This accords with diversity patterns identified in analogous Jurassic epicontinental palaeonvironments. The stratigraphic distribution of some trace fossil association boundaries accords with those of contemporary body fossil associations. However, several body fossil associations are included within the volume of distribution of a single trace fossil assemblage. □ Trace fossil association, shoreface and shelf, Oslo district, upper Ordovician.     

Ichnology of Lower Jurassic beach deposits in the Shemshak Formation at Shahmirzad,southeastern Alborz Mountains,Iran

A 19 m thick package of well-sorted lowermost Jurassic (Hettangian-Lower Sinemurian?) sandstones within the Shemshak Formation of the southeastern Alborz Mountains displays features characteristic of foreshore to upper shoreface environments such as tabular bedding, low-angle lamination, trough cross-stratification, parting lineation, and oscillation ripples. In contrast to most other beach successions recorded in the literature the sandstones contain a trace fossil assemblage characterised by low abundance but comparatively high diversity. The assemblage, comprising 14 ichnotaxa, is dominated by Palaeophycus heberti, Rhizocorallium irregulare, Gyrochorte comosa, and Parahaentzschelinia surlyki. Contrary to predictions, which assume a dominance of suspension-feeders in such high-energy environments, the trace fossil assemblage represents a variety of ethological groups ranging from suspension-feeders to deposit-feeders, detritus-feeders, scavengers, and a possible trap constructor (Ctenopholeus), whereby deposit-feeders predominate. This anomaly is explained by a high amount of organic detritus in the sediment, indicated by abundant plant material, and a position of the beach in the vicinity of a river mouth.     

Systematics and palaeoecology of middle Toarcian Asteroidea (Echinodermata) from the “Seuil du Poitou”, Western France

Complete, articulated starfish fossils are rare. However, more frequently encountered dissociated skeletal elements (ossicles) permit reliable taxonomic identification, making them a valuable data source for diversity estimates. Nearly 300 asteroid ossicles, collected from the middle Toarcian marls in western France can be assigned to five species. Four species and two genera are described: Comptoniaster vrinensis nov. sp. (Goniasteridae), Poncetaster crateri nov. gen. nov. sp. (Stauranderasteridae), Galbaster recurrans nov. gen. nov. sp. (Goniasteridae) and Pentasteria? liasica nov. sp. (Astropectinidae). The known diversity of Early Jurassic asteroids is increased from 12 to 16 species. These taxa illustrate the diversification of crown-group asteroids early in the Jurassic, following the Permo-Triassic crisis. They also reflect bias of the fossil record, and imply the existence of numerous ghost lineages in the evolutionary trees of extant groups. Variation in asteroid diversity across the “Seuil du Poitou” was driven by ecological constraints. The relative frequency (abundance of ossicles and diversity) of goniasterids and stauranderasterids increases in shallower environments. The Benthopectinidae, represented by Plesiastropecten hallovensis, occurred primarily from deep-shelf sediments. Similar ecological patterns are observed for more recent fossil and extant relatives, which further supports the idea of conservative evolution in post-Palaeozoic starfishes since the Early Jurassic.     

Chemical preservation of tail feathers from Anchiornis huxleyi,a theropod dinosaur from the Tiaojishan Formation (Upper Jurassic,China)

A panel of geochemical techniques is used here to investigate the taphonomy of fossil feathers preserved in association with the skeleton of the Jurassic theropod Anchiornis huxleyi. Extant feathers were analysed in parallel to test whether the soft tissues morphologically preserved in the fossil also exhibit a high degree of chemical preservation. Scanning electron microscopy (SEM) and energy dispersive spectroscopy (EDS) indicate that clays and iron oxide pseudomorphs occur in the surrounding sediment and also reveal the preservation of melanosome-like microbodies in the fossil. Carbon gradient along a depth profile and co-occurrence of carbon and sulphur are shown in the fossil by elastic backscattering (EBS) and particle-induced x-ray emission (PIXE), which are promising techniques for the elemental analysis of fossil soft tissues. The molecular composition of modern and fossil soft tissues was assessed from micro-attenuated total reflectance fourier transform infrared spectroscopy (micro-ATR FTIR), solid-state ¹³C nuclear magnetic resonance (CP-MAS ¹³C NMR) and pyrolysis gas chromatography mass spectrometry in the presence of TMAH (TMAH-Py-GC-MS). Results indicate that the proteinaceous material that comprises the modern feathers is not present in the fossil feathers. The fossil feathers and the embedding sediment exhibit a highly aliphatic character. However, substantial differences exist between these samples, revealing that the organic matter of the fossil feathers is, at least partially, derived from original constituents of the feathers. Our results suggest that, despite the morphological preservation of Anchiornis feathers, original proteins, that is keratin, were probably not preserved in the 160-myr-old feathers.     

A late Pleistocene progradational clastic shoreface succession in Jamaica: Implications for the preservation potential of the echinoid Leodia

A section in the Late Pleistocene Port Morant Formation of Jamaica shows a progradational deltaic succession passing from offshore muddy sands, through shoreface to foreshore deposits. The transition zone/lower shoreface has abundant Ophiomorpha burrows and contains common examples of the echinoid Leodia cf. sexiesperforata (Leske). Modern Leodia occur in shallow marine carbonate sands, and the Port Morant occurrence is the first record of abundant Leodia in the geological record. Despite their strong test construction that should give Leodia a high preservation potential, these echinoids have a poor fossil record because the environments in which they lived are rarely preserved. Consequently, the fossil record is biased towards species that occur in environments with high preservation potential.     

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.     

新疆准噶尔盆地早侏罗世掌鳞杉科植物化石 及古大气CO2浓度

植物化石气孔参数分析是目前恢复古大气二氧化碳浓度较为精准的方法之一,银杏类和松柏类等是恢复古大气CO_2浓度常用的化石类群。本文利用新疆准噶尔盆地下侏罗统三工河组的松柏类掌鳞杉科Brachyphyllum(Hirmeriella?)sp.化石对早侏罗世大气CO_2浓度进行了重建,获得早侏罗世大气CO_2浓度为～1200ppm,丰富了早侏罗世大气CO_2浓度信息,进一步说明掌鳞杉科植物通过气孔比率法在重建侏罗纪大气CO_2浓度方面的可靠性。掌鳞杉科植物的旱生构造和较高的大气CO_2浓度表明早侏罗世Toarcian期大洋缺氧事件在陆地生态系统内可能产生了一定的响应。     

New occurrences of the wood Protocupressinoxylon purbeckensis Francis: implications for terrestrial biomes in southwestern Europe at the Jurassic/Cretaceous boundary

Abstract:  Previously known from the Kimmeridgian–Portlandian of Dorset (UK) only, Protocupressinoxylon purbeckensis wood is reported here from the Kimmeridgian of Asturias (Spain) and Ajoie (Switzerland). The morphospecies taxonomy and nomenclature are discussed, and new supplementary illustrations are given. The P. purbeckensis tree was growing in dry strongly seasonal (tropophilous) environments, and the new occurrences suggest that such a climate prevailed on land all over southwestern Europe at the end of the Jurassic (Kimmeridgian sensu anglico – Portlandian). The review of fossil wood data indicates that such a stressful environment may have constrained terrestrial biocoenoses and their evolution at the Jurassic/Cretaceous boundary. But wood generic diversity curves are also strikingly similar to that drawn 20 years ago for nonmarine tetrapods, implying a fossil Lagerstätte effect.     

西北地区潮水盆地侏罗系青土井群轮藻化石

潮水盆地青土井群为一套含煤碎屑岩,在盆地东部潮参l井青土井群三段发育大量以Aclistochara占优势的轮藻化石群,其组合特征显示浓厚的晚侏罗世色彩。青土井群一、二段轮藻化石稀少,结合地层层序,推测其地质时代为早侏罗世。围岩沉积特征表明本井轮藻化石群为浅水生物群。     

Lower Jurassic Arthropod Resting Trace from the Hartford Basin of Massachusetts,USA

Cheliceratichnus lockleyi ichnogen. nov. et ichnosp. nov. is a new ichnotaxon of arthropod resting trace (cubichnium) from the Lower Jurassic (Hettangian) East Berlin Formation in Holyoke, Massachusetts, USA. The trace fossil is preserved as showing many of the external anatomical features of the exoskeleton, which resemble those of some chelicerates, notably sun spiders (Solifugae). The resting trace is directly associated with a trackway of the ichnospecies Acanthichnus cursorius Hitchcock. This is the first described fossil resting trace of a solifugan-like arthropod, and the first direct evidence of a trackmaker of A. cursorius.     

Lower Jurassic Foraminifera and Calcified Microflora from Gibraltar, Western Mediterranean

Benthic foraminifera are described for the first time from the Gibraltar Limestone Formation of the Rock of Gibraltar. The new species Siphovalvulina colomiS. gibraltarensisRiyadhella praeregularis occur with Duotaxis metula Kristan, Everticyclammina praevirguliana Fugagnoli, Siphovalvulina sp.,an atypically early example of Textulariopsis sp., and Nodosaria sp. Microflora are present as the probable cyanobacterium Cayeuxia ?piae Frollo, the alga Palaeodasycladus ?mediterraneus (Pia), and the disputed alga Thaumatoporella ?parvovesiculifera (Raineri). The foraminifera compare most closely with poorly‐known taxa from Italy, Spain and Morocco, and are consistent with an Early Jurassic (Sinemurian) age for the upper part of the &62;460‐m‐thick Gibraltar Limestone. Most are textulariids and more primitive than species well known from the later Early Jurassic (Pliensbachian) of the Mediterranean region, especially Morocco and Italy. The biota as a whole is characteristic of inner carbonate platform environments widespread along the rifted western margins of the Early Jurassic Tethys, notably those recorded from Morocco, Italy and Greece as well as southern Spain.     

Non‐traumatic bone infection in stegosaurs from Como Bluff,Wyoming

Palaeopathologies are relatively common in the fossil record; however, bone histological analyses of these are rare as they are more commonly discovered and described using radiological scans. Here, we describe the bone microstructure of infectious abscesses (osteomyelitis) in the long bones of two Stegosaurus individuals from the Como Bluff area of the Late Jurassic Morrison Formation. Multiple lytic, geographical lesions filled with reactive lamellar bone occur in a femur of a juvenile specimen. A single lesion was found in an adult tibia. This lesion is characterized by the presence of spiculated periosteal reactive growth perpendicular to the bone surface and the formation of new trabeculae within the lesion.     

Taphonomy of fossil macro-invertebrate assemblages as a tool for ecostratigraphic interpretation in Upper Jurassic shelf deposits (Prebetic Zone, southern Spain)

Composition and taphonomy of macro-invertebrate fossil assemblages, together with facies analysis, have been approached in order to interpret shifting paleoenvironmental conditions in the External Prebetic (S-SE Spain) during the early Late Jurassic (Middle Oxfordian). In oolitic and spongiolitic limestones, the size of fossil remains, mode of preservation, within-bed position, corrasion, fragmentation, epibiont and biogenic encrustation, disarticulation and uncoupling, allow recognition of two taphofacies, respectively. Identified ecostratigraphic events and trends accord with rapid flooding under high-energy conditions related to ecospace enlargement for cephalopods and then the persistence of lower energy, long-lasting exposure of skeletals and higher sedimentary rates. The paleoenvironmental interpretation is consistent with neritic environments shifting from shallow carbonate to hemipelagic sedimentation and enlarging of shelf ecospace for marine invertebrates.     

Study on the Fossil Woods Found in Zigong,Sichuan, China

This paper describes three fossil woods, which were firstly found in Zigong city, Sichuan Province, China. The localities of the fossil woods are close to the locality of the famous Dashanpu dinosaur quarry of Zigong. The strata of the fossil woods beating are Xiashaximiao Formation and Xintiangou Formation of the Middle Jurassic, approximately match the strata of the Dashanpu dinosaurs. According to the structures of the secondary xylem, the fossil woods belonged to the primitive conifer genus-Araucarioxylon. Because it was comparatively and obviously different from the other species in this genus, it is assigned as a new species: A. zigongensis. As there was neither fossil leaf nor fossil pollen or spore found in the dinosaur bearing strata before, the discovery of these fossil woods might bear some important significance in the dinosaur research.     

Crayfish burrows from Late Jurassic–Late Cretaceous continental deposits of Patagonia: Argentina. Their palaeoecological,palaeoclimatic and palaeobiogeographical significance

The trace fossil, Loloichnus baqueroensis igen. and isp. nov., from Late Jurassic–Late Cretaceous continental deposits of Patagonia, Argentina, includes thickly lined, mostly passively-filled, and Y-branched burrows. Other important features of this ichnofossil are the inner surface texture of lining showing transversal, elongated, and adjacent grooves, and less commonly, the pelletal filling of burrows. L. baqueroensis is recorded from the Bajo Grande, Bajo Tigre, Punta del Barco, and Laguna Palacios Formations, which were deposited in different volcaniclastic environments of the Deseado Massif and San Jorge Basin geological provinces, respectively. The described burrows are found in many levels of palaeosols developed in reworked piroclastic deposits, where they are the main component of the ichnofabrics, in association with meniscate and thinly lined burrows and a diffuse and complex boxwork of small diameter burrows. Root traces are also present, and in many cases, occur inside L. baqueroensis. Considering general morphology, surface texture, filling types, palaeoenvironments in which they occur, and comparisons with extant and fossil decapod burrows, the likely trace makers of L. baqueroensis were crayfishes (Decapoda: Astacidea), probably Parastacidae. The producers of L. baqueroensis inhabited soils, where their burrows probably reached the water table, and contained roots that were used for feeding. Considering climatic preferences of extant parastacids, it is proposed a temperate climate for central Patagonia during the deposition of the studied units. The widespread presence of crayfishes during Late Jurassic–Late Cretaceous times in central Patagonia, supports the monophyletic origin of this group during the Triassic, and suggests that the present restricted geographic distribution in southern South America is a relict of a broader one.     

The identification of fossil angiosperm pollen and its bearing on the time and place of the origin of angiosperms

Studies in the 1970's reporting the occurrence of fossil pollen types in the Cretaceous, coupled with surveys of extant pollen morphology of primitive flowering plants, laid the foundation for proposing a Lower Cretaceous origin of angiosperms. Over the last 30 years, morphological, ultrastructural, and ontogenetic studies of both extant and fossil pollen have provided an array of new characters, as well as greater resolution in defining character polarities. Moreover, a range of fossil pollen types exhibiting angiosperm characters occur in low frequency within Triassic and Jurassic sediments. The pollen data provide evidence of a pre-Cretaceous origin of angiosperms. Speciation and extinction rates were likely equal during the Triassic and Jurassic, resulting in the paucity of angiosperm pollen types from different geographic areas in the Atlantic – South American/African rift zone. It was not until the Lower Cretaceous that origination rates exceed extinction rates, resulting in the subsequent diversification of angiosperms and the origin of the eudicots.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

The earliest fossil evidence of bone boring by terrestrial invertebrates,examples from China and South Africa

We report the oldest fossil evidence of osteophagia by terrestrial invertebrates on both the Asian and African continents. Bones attributable to the Middle Jurassic dinosaur Chuanjiesaurus (Dinosauria: Sauropoda) were found with post-mortem insect modification in the Chuanjie Formation, Yunnan Province, China. The morphology of the borings closely matches the ichnogenus Cubiculum. Based on the lack of bioglyphs observed in Cubiculum ornatus, a new ichnospecies is proposed here. The new trace fossil, Cubiculum inornatus isp. nov., is interpreted to have been constructed for pupation by an unknown taxon of insect. Additionally, we report even older borings from Early Jurassic dinosaur bones of the Elliott Formation in the Karoo Basin, which represent the second oldest occurrence of insect traces in bone from continental settings. Both trace fossils sites have palaeogeographic implications for the origins and dispersal of osteophagia amongst terrestrial invertebrates during the Mesozoic. These discoveries push back the antiquity of pupation in animal bones by more than 100 million years to the Middle Jurassic, indicating that this behaviour, and osteophagy more generally, originated early in the Mesozoic, roughly comparable with the origination of insect pupation in woody substrates (Late Triassic).     

A new look at the Late Jurassic Canjuers conservation Lagerstätte (Tithonian,Var, France)

The Canjuers conservation Lagerstätte represents a Late Jurassic lagoonal environment. The sedimentology and stratigraphy of the locality show three different depositional sequences. Fossils are mainly found in the basalmost layers that correspond to the first phase of deposition in the lithographic limestones sensu stricto. The fossil biodiversity is rich. So far, more than 1000 specimens including 38 invertebrate and 18 vertebrate taxa have been recovered from the limestones. The depositional information suggests that most invertebrates and vertebrates were not autochthonous to the lagoon, but swept in during storm events from the open sea or nearby emerged reef environments.