Skolithos pipe rock and associated ichnofabrics from the southern Rocky Mountains,Canada: colonization trends and environmental controls in an early Cambrian sand‐sheet complex

Desjardins, P.R., Mángano, M.G., Buatois, L.A. & Pratt, B.R. 2010: Skolithos pipe rock and associated ichnofabrics from the southern Rocky Mountains, Canada: colonization trends and environmental controls in an early Cambrian sand‐sheet complex. Lethaia, 10.1111/j.1502‐3931.2009.00214.x The Lower Cambrian Gog Group of the southern Rocky Mountains of western Canada offers an opportunity to explore animal–sediment relationships in a high‐energy setting, during the early phase of Phanerozoic diversification. Its strata record a sand‐sheet complex on the broad pericontinental shelf of West Laurentia. Six ichnofabrics are recognized: Skolithos IF‐1 characterized by Skolithos linearis in planar cross‐stratified sandstone; Skolithos IF‐2 with Skolithos linearis and accessory Diplocraterion parallelum in ripple cross‐laminated, and planar and trough cross‐stratified sandstone; Skolithos IF‐3 with S. linearis and Planolites montanus in wavy‐ and flaser‐bedded sandstone, interfingering with bioturbated mudstone exhibiting P. montanus and Teichichnus rectus; Skolithos IF‐4, characterized by abundant Skolithos linearis in planar and trough cross‐stratified sandstone; Rosselia IF‐1 consisting of Rosselia isp. in planar cross‐stratified sandstone; and Rosselia IF‐2 consisting of scattered small Rosselia isp. in hummocky, cross‐stratified sandstone. The presence of contrasting ichnofabrics within a single early Cambrian sand‐sheet complex illuminates how the colonization trends of suspension and detritus feeders were controlled by factors specific to the various sub‐environments. □Gog Group, Ichnofabric, Lower Cambrian, Pipe Rock, Rosselia, Skolithos.     

山西兴县中奥陶统马家沟组五5亚段遗迹化石组合及其古环境意义

鄂尔多斯盆地东缘关家崖剖面奥陶系马家沟组五5亚段地层中保存大量无脊椎动物遗迹化石,可识别出居住迹、牧食迹、觅食迹三大类,共7个遗迹属、14个遗迹种,分别是Arenicolites isp.,Cylindricumisp.,Helminthopsis abeli,Helminthopsis isp.1,Helminthopsis isp.2,Lorenziniaisp.,Planolites isp.,Planolites beverleyensis,Planolites montanus,Teichichnus rectus,Thalassinoides isp.1,Thalassinoides isp.2,Thalassinoides isp.3,Thalassinoides isp.4。根据遗迹化石特征及其沉积环境,可划分3个遗迹组合:1、Helminthopsis-Planolites组合,该组合以觅食迹为主,部分为居住迹,出现在潮间带和潮下带;2、Thalassinoides-Teichichnus组合,该组合遗迹化石种类较单一,主要为觅食迹,分布于潮间带和潮上带;3、Thalassinoides-Helminthopsis组合,该组合出现在潮间带。基于遗迹化石组合的分布特点,提出山西兴县奥陶系马家沟组五5亚段的沉积环境模式。     

天山和昆仑—秦岭造山带志留系、石炭系和三叠系复理石相遗迹化石

在天山志留系,石炭系和东昆仑－西秦岭三叠系浊积岩系中遗迹化石发育,经研究计有：Helminthopsis hieroglyphica,Neonerietes biserialis.,N,uniserialis,palaeophycus isp.,P.serratus,P.striatus.P.tubularis,Paleodictyon imperfectum,Phycodes circinnatum,Phycosiphon incertum,Planolites annularis,P.beverleyensis,P.Montanus,Rhizocorallium ips.,Scalarituba missourensis,Skolithos ips.,?Spirorhaphe isp.,sublorenzinia ips.Tambia isp.,Taphrhelminthopsis auricularis,Teichichnus isp.,Thalassinoides isp.它们构成4种遗迹化石组合：Phycosiphon-Palaeophycus组合,Neonereites-Scalarituba组合,Paleodictyon组合和Phycodes Skolithos组合,分别代表局限缺氧深海盆地浊流沉积,半深海环境中－上扇沉积,多岛洋环境的中－下扇沉积,半深海环境的中扇浊流沉积。     

Ichnosystematics of the Lower Permian Invertebrate Traces from the Collio and Mt. Luco Basins (North Italy)

The Lower Permian ichnofauna in the Collio Formation (Artinskian) in the Val Trompia (Brescian Prealps, North Italy) has been studied for a long time, but the studies have focused mainly on vertebrate prints. In this study, the invertebrate ichnofauna of the Collio Formation, and in the epiclastites of the Monte Luco Formation (Artinskian) cropping out in the Monte Luco area (Trentino Alto Adige region), is systematically analyzed for the first time. This ichnocoenosis consists of: Permichnium isp., Paleohelcura tridactyla, Diplichnites gouldi (Types A and B), Diplopodichnus biformis, Circulichnis montanus, Helminthopsis hieroglyphica, Gordia marina, Acripes cf. multiformis, Cruziana cf. problematica, Cochlichnus anguineus, Palaeophycus tubularis, Planolites isp. and ?Scoyenia isp. The ichnoassociation from the Collio Formation belongs to the Scoyenia ichnofacies, while that from the Mt. Luco Formation belongs to the Mermia ichnofacies. The latter, because of the lack of complete data, can be linked only to one of several submersion phases of the intravolcanic basin and is not referred to the complete continental Permian sequence in this zone. The composition of the ichnoassociation here analysed is similar to those of other European (especially France and Germany) and extra-European areas (especially North America and Argentina).     

Controls on trace fossil diversity in an Early Cambrian epeiric sea: new perspectives from northwest Scotland

The Lower Cambrian Eriboll Formation of northwest Scotland is renowned for the high density, low diversity trace fossils (Skolithos ichnofacies) found in its upper Pipe Rock Member. Ichnofabric analysis of the member indicates that relatively small examples of Skolithos terminating at the same foreset boundary were formed during a brief colonization window after a single depositional event, that particularly long Skolithos specimens are equilibrichnia, and that palimpsests of Skolithos represent the marginal, slightly deeper water fringes of the Pipe Rock Member depositional environment. Nearest neighbour analysis, however, suggests that such palimpsests were uncommon. A much more diverse trace fossil assemblage is present in the overlying Fucoid Member (An t‐Sròn Formation), comprising Cruziana barbata, Dactylophycus, Didymaulichnus, Halopoa imbricata, ?Margaritichnus, Monocraterion, Monomorphichnus, Palaeophycus striatus, P. tubularis, ?Phycodes, Planolites montanus, ?Polarichnus, Rusophycus ramellensis, ?Psammichnites, Skolithos and various unidentified traces, and represents the Cruziana ichnofacies. Above the Fucoid Member, the Salterella Grit Member ichnofauna is more impoverished, yielding only Cruziana, Monocraterion, Rusophycus, Skolithos and ?Spirophyton. The ichnological variations between the Pipe Rock, Fucoid and Salterella Grit members are interpreted as being driven by changes in sea level. The low trace fossil diversity in the Pipe Rock Member indicates opportunistic colonization of laterally extensive, shoreface sediments deposited by regular influxes of terrigenous material, which were overlain by more distal, ichnologically diverse sediments (Fucoid Member) as sea level rose. A minor regression then caused an increase in terrigenous sediment input, producing an impoverished, proximal Cruziana ichnofacies (Salterella Grit Member).     

Upper Triassic (Keuper) non-marine trace fossils from the Haßberge area (Franconia, south-eastern Germany)

Various trace fossils from the Hassberge Formation and the Löwenstein Formation (Middle Keuper, Upper Triassic) of the Haßberge region are described. Twenty-three different forms have been identified, 17 of which are named, includingCruziana pascens n. isp.,Lockeia cunctator n. isp., andRusophycus versans n. isp.Lockeia siliquaria James, 1879,L. amygdaloides (Seilacher, 1953),L. triangulichnus Kim, 1994, andL. elongata (Yang, 1984) are revised and synonymized under the oldest available name,L. siliquaria James, 1879.Rusophycus eutendorfensis (Linck, 1942) andR. carbonarius Dawson, 1864 are revised. The diagnosis ofPolykladichnus Fürsich, 1981 is emended, and a diagnosis forHelminthoidichnites Fitch, 1850 is given for the first time. Among the described ichnotaxa,Skolithos ispp.,Rusophycus carbonarius, andTaenidium barretti are the most common forms. The trace fossil association is typical of theScoyenia ichnofacies, which indicates non-marine, periodically or completely inundated environments, such as floodplains and lake margins. Two palaeoichnocoenoses are identified. One ichnocoenosis, dominated byCruziana problematica, cf.Polykladichnus isp., andSkolithos isp. B characterizes margins of trough cross-bedded sandstones. Another ichnocoenosis, dominated byRusophycus versans n. isp.,Taenidium barretti,Scoyenia gracilis andSkolithos isp. A is related to ephemeral lake deposits. Taxonomic recommendations for the use of hitherto described and figured invertebrate Keuper trace fossils from Germany are given.     

A diverse deep‐marine Ichnofauna from the Eocene Tarcau sandstone of the Eastern Carpathians,Romania

The Paleocene to Middle Eocene Tarcau Sandstone at Buzau Valley, eastern Carpathians, Romania, records sedimentation in a turbidite system. These strata contain a diverse and abundant pre‐ and postdepositional ichnofauna consisting of 35 ichnogenera and 54 ichnospecies. The predepositional assemblage is rich in graphoglyptids and ornate grazing trails; simple grazing trails, resting traces, and feeding structures also occur. The predepositional assemblage includes Acan‐thorhaphe, Belorhaphe, Cardioichnus, Circulichnus, Coch‐lichnus, Cosmorhaphe, Desmograpton, Fustiglyphus, Gordia, Helicolithus, Helminthopsis, Helminthorhaphe, Lorenzinia, Megagrapton, Paleodictyon, Paleomeandron, Protopaleodictyon, Scolicia (S. strozzii), Spirorhaphe, Spirophycus, Treptichnus, and Urohelminthoida. The ich‐nodiversity, composition, ethology, and morphologic complexity of the predepositional association are indicative of the Nereites ichnofacies. The postdepositional association essentially consists of dwelling, feeding, and grazing traces, and is represented by Chondrites, Glockerichnus, Halopoa, Nereites, Ophiomorpha, Phycodes, Planolites, Polykampton, Scolicia(S. prisca. S. striata), Taenidium, Thalassinoides, and Zoophycos. Palaeophycus occurs in both assemblages. Allochthonous Teredolites is present in wood fragments, The postdepositional association includes elements of the Skolithos ichnofacies and facies‐crossing forms that are commonly present in deep‐marine deposits, Elements of the Skolithos ichnofacies are present not only in the most proximal parts of the turbidite system, but also in distal parts. The number of predepositional forms greatly exceeds postdepositional ones, reflecting a dominance of K‐selected over r‐selected population strategies in a stable environment. High levels of ichnodiversity in the Tarcau Sandstone are comparable with deep‐sea ichnofaunas from the Polish Carpathians and with other flysch trace‐fossil assemblages of similar age. This abundant and diverse Eocene ichnofauna supports the idea of extremely rich deep‐sea ichnofaunas in the Cenozoic.     

Devonian Trace Fossils of the Horlick Formation,Ohio Range,Antarctica: Systematic Description and Palaeoenvironmental Interpretation

The clastic Horlick Formation contains an ichnofauna of 28 ichnotaxa dominated by burrowers. These are a mixture of simple vertical forms (Skolithos linearis, S. magnus, Bergaueria cf. langi, Rosselia socialis, Monocraterion isp.), U or arc-like forms (Diplocraterion parallelum, Arenicolites types A and B, Catenarichnus antarcticus, C. isp., aff. Lanicoidichna isp.), and complex, vertical spiral structures (Asterosoma isp., Spirophyton isp.). Horizontal burrows include Ancorichnus cf. capronus, Palaeophycus tubularis, and Psammichnites devonicus isp. nov. Surface traces comprise Haplotichnus isp., Cruziana problematica, C. rhenana, Rusophycus aff. carbonarius, R. isp., Protovirgularia rugosa, Lockeia ornata and cubichnia indet., while trackways include Diplichnites gouldi, D. isp., Maculichna? isp. and large imprints. The Horlick Formation (maximum 56 m) records an early Devonian transgression onto a deeply weathered land area that lay in the direction of Marie Byrd Land, spreading from the South Africa sector of Gondwana. Analysis of the trace fossils confirms their formation in near-shore to intertidal environments, with some ichnotaxa living close to the marine/fluvial boundary (e.g., Spirophyton, aff. Lanicoidichna, Cruziana problematica, Rusophycus aff. carbonarius, Maculichna?). The ichnofauna probably comprised a variety of marine suspension and deposit-feeding worms, shallow burrowing molluscs (bivalves and possibly bellerophontids) and several different types of arthropods, including trilobites.     

Environmental variability of Macaronichnus ichnofabrics in Eocene tidal‐embayment deposits of southern Patagonia,Argentina

The Middle Eocene Man Aike Formation of southern Patagonia, Argentina, offers the opportunity to study the palaeoecological and palaeoenvironmental significance of Macaronichnus ichnofabrics in a tide‐dominated, coastal embayment compound‐dune complex. Four recurrent Macaronichnus ichnofabrics are recognized: Macaronichnus IF‐1 characterized by Macaronichnus segregatis segregatis; Macaronichnus IF‐2 with M. segregatis segregatis, Ophiomorpha nodosa, Skolithos linearis and rare Palaeophycus tubularis; Macaronichnus IF‐3 typified by M. segregatis segregatis, O. nodosa, Asterosoma radiciforme and rare S. linearis; and Macaronichnus IF‐4 consisting of M. segregatis segregatis, Nereites missouriensis, A. radiciforme, O. nodosa, Rosselia socialis, S. linearis, P. tubularis, Planolites isp. and Chondrites isp. Variations in these ichnofabrics are linked to changes in local hydraulic conditions in the upper and lower trough areas of the compound dunes. In high‐energy, shallow and marginal marine environments, the Macaronichnus tracemaker is commonly the first to colonize the shifting, sandy substrates. Overprinting by other ichnoguilds occurs only after the physiochemical conditions became favourable and the colonization window length increases.     

山东省东北部车镇凹陷古近系沙河街组滨浅湖沉积中的遗迹群落

济阳坳陷车镇凹陷古近系沙河街组滨浅湖沉积中产有十分丰富的动物遗迹化石,共鉴定出遗迹属 10个,未定属 1个,其中遗迹种 7个,未定种 4个。遗迹化石包括Beaconitesantarcticus, Beaconitescapronus, Planolitesmontanus, Macaronichnussegregatis, Taenidiumbarretti, Cochlichnusanguineus, Gordiasp., Skolithossp., Ophiomorphanodosa, Sagittichnussp.和Favreinasp.。这一遗迹化石群落主要是无脊椎动物的进食迹、觅食迹、居住迹和停息迹,其中大部分呈全浮痕保存,少数呈上浮痕保存,并形成于经常干旱和周期性泛滥的滨浅湖沉积环境中。该群落可识别出两个遗迹化石组合,即 1)Beaconites Taenidium组合,它主要产自干旱气候条件下的滨浅湖沉积环境;2)Sko lithos Ophiomorpha组合,它代表了丰水期水体能量相对较高的滨湖沉积环境。     

A Nereites ichnofacies from the Ordovician‐Silurian Welsh Basin

The sediments of the Ordovician‐Silurian succession of west Wales are interpreted as turbiditic sands with associated pelagic and hemi‐pelagic mudstones. The strata contain a trace fossil assemblage consisting of fifteen ichnogenera, namely: Chondrites, Circulichnis, Cochlichnus, Cosmorhaphe, Desmograpton, Gordia, Helminthoida, Helminthopsis, Neonereites, Nereites, Paleodictyon, Palaeophycus, Planolites, Protopaleodictyon and Spirophycus. The ichnofaunal assemblage is typical of the Nereites ichnofacies. Twenty three ichno‐species are described, two of which (Helminthoida granulata, Palaeophycus serratus) are new. Turbiditic sands contain the most abundant and diverse ichnoassemblages, a reflection of favorable environmental conditions for habitation by benthic organisms. Both the turbiditic mudstones and anoxic hemipelagites contain just two ichno‐species, namely Chondrites ichnosp. and Planolites montanus both of which are common, albeit less so in the latter facies. The effects of preservation potential, medium and preservation mechanisms are also important controls on distribution.     

Trace fossils from arenig flysch sediments of eire and their bearing on the early colonisation of the deep seas

A sequence of Lower Ordovician (Arenig) turbidites in Co. Wexford, Eire, has yielded one of the earliest diverse ichnofaunas yet recorded from deep water sediments comprising: Chondrites, Glockerichnus, Gordia, Helminthopsis, Lorenzinia, Neonereites, Palaeophycus, Paleodictyon, Planolites, Sublorenzinia, Taenidium, Taphrhelminthopsis, Teichichnus and Tomaculum. This ichnofauna is critical in any analysis of the colonisation of the deep seas by trace fossil‐producing animals.

A world‐wide review shows that the earliest trace fossils are mainly from Late Precambrian shelf sea environments, but many more evolved during very rapid diversification in the pre‐trilobite Lower Cambrian.

There was little increase in diversity in shallow water after the Lower Cambrian but a progressive colonisation of the deep ocean took place and this accelerated during the Ordovician, when the main lineages of deep sea trace fossils were established there. Rosetted, patterned, meandering and simple spiral forms evolved in shallow water in the Upper Precambrian and pre‐trilobite Lower Cambrian and only later migrated into the deep sea, whereas complex, closely programmed, spiral traces may have evolved there.     

A Late Triassic Invertebrate Ichnofauna from Ghost Ranch,New Mexico

Invertebrate trace fossils are reported from the Late Triassic Chinle Formation at Ghost Ranch, New Mexico, for the first time. They occur in beds higher in the section than the Coelophysis Quarry. Six ichnotaxa are recognized: Steinichnus milfordensis, Planolites montanus, Palaeophycus tubularis, Taenidium serpentium, ?Arenicolites sp. and Skolithos sp. This ichnofauna is consistent with the Scoyenia ichnofacies, considered typical of lake margins in semiarid to arid regions.     

Tetrapod and invertebrate trace fossils from aeolian deposits of the lower Permian of central-western Argentina

Abundant tetrapod footprints are described from the Early Permian Yacimiento Los Reyunos Formation including both collected and in situ specimens. The slabs come from several quarries at the Sierra Pintada and Sierra de las Peñas area, south-west of Mendoza, Argentina. The trace fossil assemblage, which constitutes one of the oldest known from Gondwana, comprises excellent-preserved tetrapod tracks (Chelichnusduncani, Chelichnusgigas and ‘pear-like’ footprints) and invertebrate simple sub-horizontal (Palaeophycustubularis) and vertical (Skolithos isp.) burrows formed in a aeolian dune field. The analysis of the tetrapod track producers indicates the presence of at least three different taxa of sprawling to semi-erect therapsids, thus suggesting the presence of members of this clade, or closest relatives, in the Early Permian of southern Gondwana. Moreover, a series of measurements and simple indexes were developed to estimate body proportions and locomotion styles of the putative trackmakers. The new assemblage, analysed in the context of other known Permian assemblages from Pangea, is one the few known in Gondwana to be present in an aeolian environment. The evaluation of the assemblage, in the light of aeolian ichnofacies (Chelichnus, Octopodichnus and Entradichnus), shows that it has common elements with the Chelichnus and Entradichnus ichnofacies.     

Palaeoenvironmental implications of the ichnology and geochemistry of the Westbury Formation (Rhaetian), Westbury‐on‐Severn,south‐west England

Abstract: The Westbury Formation (Rhaetian) beds of Westbury Garden Cliff, Westbury‐on‐Severn, west of Gloucester, Britain, show an unusual combination of features. Both deep water and emergent characteristics are present within the sediments and the trace fossils. The ichnoassemblage consists of abundant Selenichnites, Planolites beverlyensis and Lockeia with rarer Oniscoidichnus, Chondrites, Rhizocorallium irregulare, Taenidium serpentium, an unusual form of Walcottia and Merostomichnites‐like traces. These trace fossils display an interesting relationship with the sediments: low‐energy Cruziana ichnofacies is found within high‐energy sandstones. The sandstones are interbedded with laminated mudstones, apparently deposited in deep water, but some aspects of the ichnoassemblage, preservation and sedimentation indicate shallower water. One new trace fossil, Radichnus allingtona igen. et isp. nov., closely resembles the traces of modern fiddler crabs and imply emergence, by analogy. This ichnofauna is similar to early stage disaster colonisation in recent experiments in Long Island Sound (south of Connecticut, USA) and with storm‐influenced deposits within the Cardium Formation (Seebe, Alberta, Canada). This indicates a lagoonal environment with influxes of sand and oxygen. Total organic carbon levels were found to fluctuate greatly between stratigraphic layers but remained relatively high. This implies low oxygen conditions. The abundance of sulphur (in pyrite) also supports an interpretation of anoxic conditions, and low sedimentation rates within the shale layers. A restricted shallow basin or lagoonal environment is proposed for the palaeoenvironment, with fluctuating oxygen influencing diversity.     

Trace fossils analysis of fluvial to open marine transitional sediments: Example from the Upper Devonian (Geirud Formation), Central Alborz,Iran

This study integrates ichnological and sedimentological data to interpret depositional environments of the mixed siliciclastic-carbonate fluvial to marine sediments of the Geirud Formation (Upper Devonian) in the central Alborz, northern Iran. Lithofacies analysis shows that these sediments are deposited in fluvial, tidal, shoreface, and shelf environments. Fluvial and tidal deposits are characterized by the presence of bi- to multi-directional cross beddings, reverse directional current ripples, low angle cross beddings, and herringbone cross beddings, with a few scattered Skolithos and Palaeophycus. Shoreface sediments, accumulated in a storm-influenced setting, are characterized by a preferentially interface and low diversity Cruziana ichnoassemblage (Rhizocorallium, Thalassinoides, Palaeophycus, Chondrites, and Ophiomorpha). In contrast to the fluvial-tidal assemblage, the storm-influenced shelf sediments display a highly diversified, mixture of dwelling and feeding forms (Arenicolites, Protovirgularia, Diplocraterion, Palaeophycus, Thalassinoides, Chondrites, and Helminthopsis), reflecting the presence of adequate food resources both in substrate and water column under normal salinity conditions. A fluvial-shelf replacement of the weakly to scarcely bioturbated sediments by the Rhizocorallium-Thalassinoides suite (Cruziana) – Chondrites-Helminthopsis (distal Cruziana) suite from the lower to upper parts of the succession clearly indicates an overall deepening upward in the Geirud Formation. In contrast to the lower part, generally of restricted environment, the upper part of the succession mainly shows open marine conditions. Ichnofabric development is controlled primarily by depositional conditions, e.g., bottom water oxygenation, sediment type, food abundance, and hydrodynamic level, which all exert control on substrate colonization style.     

An integrative ichnological and taphonomic approach in a transgressive–regressive cycle: a case study from Devonian of Paraná Basin,Brazil

The palaeoenvironmental context of a section of the Devonian Ponta Grossa Formation (Paraná Basin) was examined using an integrated ichnological and taphonomic approach. Three taphofacies (T‐A, T‐B and T‐C) and six ichnofabrics are recognized. T‐A is mainly composed of disarticulated organisms and is associated with Chondrites, Planolites‐Palaeophycus, Asterosoma‐Zoophycos and Asterosoma‐Chondrites ichnofabrics, representing the Cruziana ichnofacies. The upward increase in diversity and abundance of body fossils for Taphofacies A can be explained by changes in sedimentation rates during aggradational to progradational phases. T‐B beds, represented by articulated brachiopods, occur in distal tempestites associated with the Skolithos ichnofabric (Skolithos ichnofacies) and represent higher hydrodynamic energy and oxygenation than in T‐A. Taphofacies C, characterized by a mix of articulated and disarticulated organisms, generally lacks bioturbation except for Chondrites ichnofabrics occurring only at the bed tops, suggesting low substrate oxygenation and low‐energy conditions. T‐C indicates the maximum transgression in the T‐R cycle, characterizing it as a good indicator of maximum flooding surfaces. The composite Asterosoma‐Teichichnus ichnofabric (bioturbation degree 5) occurs in layers without body fossils and represents suites of the Cruziana ichnofacies. This likely reflects intense intrastratal activity at all substrate levels, which facilitated the oxygenation of the substrate disallowing the preservation of organism remains. This observation indicates that the bioturbation is an important factor controlling the loss of taphonomic information within the TAZ, mainly when associated with detritus‐feeding trace fossils.     

Lopingian (Upper Permian) trace fossils from the northern Penglaitan Section,Laibin, Guangxi,South China and their environmental implications

The end-Permian mass extinction has been widely documented to be accompanied with oceanic anoxia, which was considered as one of the most plausible killing mechanisms. However, it is still unclear when anoxia began to occur and how widely affected during the pre-extinction interval. In this study, Lopingian bottom-water oxygen level changes around the Wuchiapingian–Changhsingian boundary (WCB) at the northern Penglaitan section in Guangxi are analysed based on trace fossil assemblages and ichnofabric indices. Detailed bed-by-bed ichnological analyses confirm the presence of trace fossils Chondrites intricatus, C. isp., Palaeophycus isp., Planolites isp. A, P. isp. B, Thalassinoides isp. A, T. isp. B, Trichichnus linearis, Zoophycos isp., and other bioturbational structures. The ichnofabric indices around the WCB are mostly 1–2, but rapidly increase to 4–5 in some short intervals. The distinct fluctuations of ichnofabric indices suggest that the oxygen level of sediments was generally deficient, but the dysoxic or anoxic periods were punctuated by several short oxic intervals. Thus, it can be inferred that the benthic organisms were under secularly environmental stresses around the WCB at the northern Penglaitan section. The anoxic conditions unfavourable for aerobiosis in the deep basin may start in the late Wuchiapingian. However, more sections need to be studied to determine whether this is a local or regional phenomenon.     

Bored and Burrowed: An Unusual Echinoid Steinkern from the Type Maastrichtian (Upper Cretaceous,Belgium)

A test of the irregular echinoid Hemipneustes striatoradiatus (Leske) was dissolved in dilute hydrochloric acid to expose the flint steinkern. This revealed slender, string-like borings that were mainly limited to ambulacra (Talpina isp. cf. T. canna (Price)); less common pouch-like borings limited to part of one ambulacrum only (Rogerella isp. cf. R. pattei (Saint Seine)); and slender, sinuous to contorted burrows, Planolites montanus Richter. Ingress to the test (borers) and sediment infill (burrowers) was presumably through the ambulacral pore pairs of the dead test. Borings and burrows were probably not coeval. Such infestations of fossil echinoids may be locally common, but their identification is controlled by the difficulty of seeing features of both the test and the infill.     

Environmental distribution of trace fossils in the Jurassic of Kachchh (Western India)

Summary Trace fossils occur abundantly in Middle Jurassic rocks of the Kachchh Basin. They are found in environments ranging from beach sequences down to central parts of the basin. For stratinomic reasons, they are particularly well preserved in storm deposits. Their distribution pattern exhibits a clear relationship to the hydrodynamic conditions and, secondarily, to bathymetry, and follows the classic ichnofacies concept ofSeilacher (1967). High energy nearshore areas and submarine shoals are represented by members of the Skolithos ichnofacies such asOphiomorpha nodosa, Arenicolites, Diplocraterion parallelum, andRhizocorallium jenense. The storm-influenced ramp contains both members of the Cruziana ichnofacies (e.g.Rhizocorallium irregulare, Thalassinoides suevicus, Taenidium serpentinum, Chondrites) and the Skolithos ichnofacies (in particularOphiomorpha). The former were produced during interstorm phases, the latter are of post-storm origin. Carbonate ramp environments of low to intermediate energy also contain members of the Cruziana ichnofacies, whilst equivalent siliciclastic environments are characterized by a low-diversity Zoophycos ichnofacies. Low energy basinal environments of fine-grained substrates contain an impoverished Cruziana ichnofacies consisting ofChondrites, Trichichnus andThalassinoides suevicus. 32 ichnotaxa are briefly described, among themSphaerichnus lobatus ichnogen. et ichnosp. nov.