A systematic overview of fossil osmundalean ferns in China: Diversity variation,distribution pattern,and evolutionary implications

The order Osmundales is a unique fern taxon with extensive fossil records in geological past. Diverse osmundalean fossils have been reported from China, ranging in age from the Late Palaeozoic to the Cenozoic. Most of them are based on leaf impressions/compressions, but permineralized rhizomes are also well documented. In this study, we provide a systematic overview on fossil osmundalean ferns in China with special references on diversity variations, distribution patterns, and evolutionary implications. Fossil evidence indicates that this fern lineage first appeared in the Late Palaeozoic in China. The Late Triassic to Middle Jurassic interval was the radiation stage. From the Late Jurassic onward, fossil diversity declined rapidly. Cenozoic osmundalean taxa are represented by the relict species of Osmunda. Geographically, osmundalean fossils are found from both the Northern and Southern phytoprovinces of China, though variations are documented for geographical ranges. The Chinese fossil records cover almost all important stages for the macroevolution of the Osmundales, and contribute to further understanding of evolutionary processes of this peculiar fern lineage.     

遗迹化石Chondrites的指相意义和阶层分布

介绍和评述Chondrites的基本特征和流行的成因解释 ,记述最近发现于美国犹他州中西部北美下 中奥陶统之交层型剖面上的Chondrites遗迹组构的不寻常特征和属性 :细小的Chondrites被所有与之共生的遗迹化石交切 ,形成于水与沉积物界面附近缺氧的早期软底质中 ,Chondrites是形成最早和最浅的遗迹阶层。流行的遗迹相和遗迹阶层模式的不足是 :前者在操作上 ,简单地将遗迹化石及其组合处理为同沉积或准同沉积的物理沉积构造 ,忽略了造迹活动的多阶段和多阶层的特点 ;后者的资料基础仅依据中生代以来的地层记录 ,对重大生物 环境事件和造迹生物生态习性演化对遗迹阶层形成和分布的影响注重不够。指出Chondrites的阶层分布和对古氧相的示踪在古生代与中生代有重要差别     

New evidence of nearshore Mid-Triassic <Emphasis Type="Italic">Zoophycos</Emphasis>: morphological and paleoenvironmental characterization

Zoophycos is a well-known trace fossil common throughout the Phanerozoic. Paleozoic forms show important differences in morphology and habitat distribution with respect to the Jurassic, Cretaceous, and Cenozoic ones. Therefore, Early–Middle Triassic is considered a crucial time-span for the understanding of the evolution of this trace fossil. So far, Early Triassic Zoophycos is unknown and Middle Triassic forms were recorded only in deposits from Thuringia. The morphology and paleoenvironment of Zoophycos from the middle–upper Muschelkalk of the Iberian Range is herein described. The best-preserved trace fossils occur in a dolomicritic bed Ladinian in age, and are represented by small forms with a subcircular, slightly lobed outline and very little penetration depth. They were deposited in a very shallow, quiet-water environment with transition to supratidal/emerged areas. The low diversity of both trace fossils and skeletal remains point to stressful conditions related to strong salinity variations and/or poor water circulation. A comparison was made with Zoophycos from Anisian deposits of the Muschelkalk in Germany. This showed that both forms are quite simple and penetrate only the shallowest tiers, although they are different in whorl outline and lobe shape. This confirms that, notwithstanding the morphological variability of this group, Zoophycos still maintained a quite simple structure in the Triassic. A shallow-water environment was deduced for both localities, confirming that at least until the Early Jurassic Zoophycos had not definitively migrated toward deep-water areas.     

A Lower Cambrian shallow-water occurrence of the branching ‘deep-water’ type trace fossilDendrorhaphe from the Lontova Formation, eastern Latvia

We describe a unique Lower Cambrian (Lontova Formation, Latvia) occurrence of a branching trace fossil assigned toDendrorhaphe isp. In the Mesozoic and Cenozoic this type occurs exclusively in deep-water flysch deposits.Dendrorhaphe may, therefore, be an additional example of a flysch-type trace fossil with an origin in early Palaeozoic shallow-water environments and subsequent onshore-offshore displacement, or perhaps, expansion. Recently,Dendrorhaphe Seilacher 1977, has been synonymized withDendrotichnium Farrés 1967 andUbinia Grosshejm 1961. This needs further consideration, especially because of possible relation of these ichnogenera toChondrites patulus Fischer-Ooster 1858.     

First occurrence of the Ichnogenus Selenichnites from the Middle Jurassic Strata of the Skoura Syncline (Middle Atlas,Morocco); Palaeoecological and palaeoenvironmental context

Mesozoic strata of North Africa yield the first occurrence of the ichnogenus Selenichnites. The trace fossils occur on the top surface of a sandy carbonate deposit in the axis of a Middle Atlas syncline (Skoura Syncline, NE Morocco). The ichnofossil-bearing horizon belongs to the Late Bajocian–Early Bathonian Ich Timellaline/Bou Akrabene Formation. The trace fossils are crescent-shaped and the best preserved exhibits a posterior central axial impression (possible telson tail impression). They are interpreted as feeding burrows (fodinichnia) or hiding depressions of Xiphosurids or Limulids (horseshoe crabs) on a sandy carbonate substrate beneath a veneer of muddy deposits. The sedimentological character suggests a relatively protected shallow water subtidal palaeoenvironment preceding the Bathonian regression of the Atlas domain. This discovery provides the first evidence of xiphosurans or xiphosuran-like organisms inhabiting the southern shores of the Tethys in the Middle Jurassic.     

川西南峨眉山地区上白垩统夹关组遗迹组构特征及其环境意义

川西南峨眉山地区上白垩统夹关组由一套厚层到块状中、细砂岩组成,夹有少量粉砂岩和泥岩,底部有砾岩。本组地层中发育有丰富的遗迹化石,共识别出9属11种。根据遗迹化石的分布情况,划分出了5种遗迹组构,即:Palaeophycus遗迹组构、Arenicolites遗迹组构、Skolithos遗迹组构、Planolites遗迹组构和Scoyenia遗迹组构;结合它们所分布岩层的沉积特征,认为前三者分别分布于河流环境中水动力强的边滩下部、水动力中等到较强的边滩中上部和边滩向堤岸过渡的环境,后两者分布于水动力较弱的天然堤环境。     

Global Taxonomic Diversity of Anomodonts (Tetrapoda,Therapsida) and the Terrestrial Rock Record Across the Permian-Triassic Boundary

The end-Permian biotic crisis (∼252.5 Ma) represents the most severe extinction event in Earth''s history. This paper investigates diversity patterns in Anomodontia, an extinct group of therapsid synapsids (‘mammal-like reptiles’), through time and in particular across this event. As herbivores and the dominant terrestrial tetrapods of their time, anomodonts play a central role in assessing the impact of the end-Permian extinction on terrestrial ecosystems. Taxonomic diversity analysis reveals that anomodonts experienced three distinct phases of diversification interrupted by the same number of extinctions, i.e. an end-Guadalupian, an end-Permian, and a mid-Triassic extinction. A positive correlation between the number of taxa and the number of formations per time interval shows that anomodont diversity is biased by the Permian-Triassic terrestrial rock record. Normalized diversity curves indicate that anomodont richness continuously declines from the Middle Permian to the Late Triassic, but also reveals all three extinction events. Taxonomic rates (origination and extinction) indicate that the end-Guadalupian and end-Permian extinctions were driven by increased rates of extinction as well as low origination rates. However, this pattern is not evident at the final decline of anomodont diversity during the Middle Triassic. Therefore, it remains unclear whether the Middle Triassic extinction represents a gradual or abrupt event that is unique to anomodonts or more common among terrestrial tetrapods. The end-Permian extinction represents the most distinct event in terms of decline in anomodont richness and turnover rates.     

Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

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.     

Palaeoecological and phylogenetic implications of a new scleractiniamorph genus from Permian sponge reefs,south China

Scleractinian corals are the most important constituents of modern coralgal reefs. For many years, it was thought that they first appeared in the Middle Triassic and subsequently underwent explosive radiation. However, abundant scleractinian-like corals with ancestral morphological traits have recently been recovered from Middle Permian sponge reefs in China, which not only confirms a role in Permian reef ecology but also suggests a possible Palaeozoic origin for the group. Two species of a new Permian scleractiniamorph genus from China are described herein as Houchangocyathus wangi gen. et sp. nov. and Houchangocyathus yaoi gen. et sp. nov. Putative Palaeozoic Scleractinia may have evolved over a substantial time interval and diverged into stem lineages by the end of the Permian. These forms evolved within both the rigid framework of their basic body plan and the morphological constraints characteristic of each lineage. The Middle Permian development of calcisponge reefs was closely related to habitat expansion, which would have provided an ideal dwelling for scleractinian-like corals and enhanced their chances of fossilization. Such scleractiniamorphs disappeared at the end-Permian extinction, but may have survived as progenitors of Triassic Scleractinia.     

Delayed recovery of non-marine tetrapods after the end-Permian mass extinction tracks global carbon cycle

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom-bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.     

The Trace Fossil Paleodictyon within The Cruziana Ichnofacies: First Record from The Devonian in Pennsylvania

Offshore marine deposits of hemipelagic dark-gray shales comprising the Middle Devonian Mahantango Formation have yielded the first evidence of the trace fossil Paleodictyon from Pennsylvania. Paleodictyon occurs in conjunction with a diversity of largely deposit-feeding trace fossils belonging to the Cruziana ichnofacies, and documents another example of a shallower-water occurrence of this ichnofossil in Paleozoic rocks.     

Fossil fungi with suggested affinities to the Endogonaceae from the Middle Triassic of Antarctica

Documented fossil evidence of zygomycetous fungi is rare. A conspicuous fungal fossil, Jimwhitea circumtecta gen. et sp. nov., occurs in permineralized peat from the Middle Triassic of Antarctica. The fossil is interpreted as a mantled zygosporangium that buds from a macrogametangium subtended by a sac-like macrosuspensor. The macrogametangium is united at its tip with a microgametangium which is subtended by a micro-suspensor. This configuration is strikingly similar to the zygosporangium-gametangia complexes seen in certain modern Endogonaceae. Co-occurring with J. circumtecta are isolated propagules closely resembling the zygosporangium of J. circumtecta and a portion of a sporocarp containing zygosporangia embedded in a gleba. Several of the sporangia are borne on ovoid or elongate structures, which we interpret as gametangia. These fossils offer an exceptionally detailed view of the morphology and reproductive biology of early Mesozoic zygomycetes.     

Diversity variation and tempo-spatial distribution of Otozamites (Bennettitales) in the Mesozoic of China

Otozamites is a representative fossil leaf morphogenus of the extinct Bennettitales, with an extensive distribution during the Mesozoic, especially in China. Understanding the fossil diversity variation and distribution pattern of Otozamites in China will provide information on biodiversity of bennettitalean plants as well as for reconstruction of palaeogeography and palaeoclimate conditions during the Mesozoic. So far, 46 species of this genus have been described in China, excluding unspecified species. The results show that the fossils of Otozamites are extensively recorded in the Late Triassic, and then reach their maximum development in the Early Jurassic, followed by a reduction in diversity in the Middle and Late Jurassic, and finally become extinct at the end of Early Cretaceous. Geographically, they occur in both Northern and Southern Floristic Provinces in the Mesozoic of China, with a relatively higher abundance in the Southern Floristic Province. It implies that the diversity variation and distribution of Otozamites are closely related to the change of the palaeoclimatic conditions. The warm and humid climate prevailed in the Late Triassic and Early Jurassic in South China, propitious to the development of Otozamites. After the Middle Jurassic, dry and hot climate may have caused the lower diversity level and blocked the development of Otozamites; finally at the end of the Early Cretaceous, the frequent arid climate may be a major cause for the extinction of Otozamites.     

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 stem-neopterygian fish from the Middle Triassic of China shows the earliest over-water gliding strategy of the vertebrates

Flying fishes are extraordinary aquatic vertebrates capable of gliding great distances over water by exploiting their enlarged pectoral fins and asymmetrical caudal fin. Some 50 species of extant flying fishes are classified in the Exocoetidae (Neopterygii: Teleostei), which have a fossil record no older than the Eocene. The Thoracopteridae is the only pre-Cenozoic group of non-teleosts that shows an array of features associated with the capability of over-water gliding. Until recently, however, the fossil record of the Thoracopteridae has been limited to the Upper Triassic of Austria and Italy. Here, we report the discovery of exceptionally well-preserved fossils of a new thoracopterid flying fish from the Middle Triassic of China, which represents the earliest evidence of an over-water gliding strategy in vertebrates. The results of a phylogenetic analysis resolve the Thoracopteridae as a stem-group of the Neopterygii that is more crown-ward than the Peltopleuriformes, yet more basal than the Luganoiiformes. As the first record of the Thoracopteride in Asia, this new discovery extends the geographical distribution of this group from the western to eastern rim of the Palaeotethys Ocean, providing new evidence to support the Triassic biological exchanges between Europe and southern China. Additionally, the Middle Triassic date of the new thoracopterid supports the hypothesis that the re-establishment of marine ecosystems after end-Permian mass extinction is more rapid than previously thought.     

Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschelkalk) mud flat

Knaust, D. 2010: Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschelkalk) mud flat. Lethaia, Vol. 43, pp. 344–356. A new Fossil‐Lagerstätte is reported from the Middle Triassic of Germany, preserving Foraminifera, Nematoda, Platyhelminthes, Nemertea, Annelida and a range of Arthropoda together with their traces. This is the oldest fossil record of free‐living nematodes and turbellarians, and the first occurrence of nemerteans in the Mesozoic. The rare preservation of the benthic associations together with their traces is unique; especially the abundant occurrence of different phyla as meiofauna (organisms with shortest dimension between 0.06 and 1 mm) provides an insight into the palaeoecological conditions of a 240‐Ma‐old muddy tidal flat. The preservation of benthic animals at the termination of their traces offers the exclusive opportunity to assign producers to the trace fossils. The results show that the discussed phyla were already established in the early Mesozoic in a similar diversity and composition as in modern analogues. The new Fossil‐Lagerstätte has the potential to prove a number of soft‐bodied taxa immediately after the end‐Permian mass extinction, which has wide‐ranging implications for phylogenetic interpretations. □Benthos, soft‐bodied organisms, Fossil‐Lagerstätte, Muschelkalk, trace fossils, Triassic.     

Diverse bryophyte mesofossils from the Triassic of Antarctica

Compared with the fossil record of vascular plants, bryophyte fossils are rare; this circumstance is probably related to a lower preservation potential compared with that of vascular plants. We searched for bryophyte remains in extensive collections of plant‐fossil assemblages from the Triassic of Antarctica and identified three assemblages with surprisingly well‐preserved bryophyte fossils. Although most bryophyte remains are too fragmented to conclusively place them in a detailed systematic context, they exhibit features sufficient to suggest the presence of at least four types of leafy bryophytes and two orders of thallose liverworts (Pallaviciniales and Metzgeriales) in the high‐latitude Triassic ecosystems of Antarctica. The leafy bryophytes exhibit combinations of morphological features (e.g. keeled and entire‐margined, ecostate leaves with elongated cells) that today occur in only a few small, systematically isolated groups, but were common among Palaeozoic and especially Mesozoic bryophytes. The diverse morphologies of the bryophyte fossils add further support to previous hypotheses that during warmer periods in the Earth's history, bryophyte vegetation may have been particularly rich and diverse in high‐latitude regions. Through analysis of the sedimentology and taphonomy of these assemblages, we identify a combination of key factors that may explain the preservation of bryophyte fossils in these deposits: (1) punctuated, high‐energetic sedimentary events causing traumatic removal and incorporation of bryophytes into sediment‐laden flood waters; (2) limited transport distance, and short period of suspension, followed by rapid settling and burial as a result of a rapidly decelerating flow discharging into a floodplain environment; and (3) early‐diagenetic cementation with iron hydroxides in locally anoxic zones of the organic‐rich, muddy substrate.     

Trace fossil analysis of lacustrine facies and basins

Two ichnofacies are typical of lacustrine depositional systems. The Scoyenia ichnofacies characterizes transitional terrestrial/nonmarine aquatic substrates, periodically inundated or desiccated, and therefore is commonly present in lake margin facies. The Mermia ichnofacies is associated with well oxygenated, permanent subaqueous, fine-grained substrates of hydrologically open, perennial lakes. Bathymetric zonations within the Mermia ichnofacies are complicated by the wide variability of lacustrine systems. Detected proximal–distal trends are useful within particular lake basins, but commonly difficult to extrapolate to other lakes. Other potential ichnofacies include the typically marine Skolithos ichnofacies for high-energy zones of lakes and substrate-controlled, still unnamed ichnofacies, associated to lake margin deposits. Trace fossils are useful for sedimentologic analysis of event beds. Lacustrine turbidites are characterized by low-diversity suites, reflecting colonization by opportunistic organisms after the turbidite event. Underflow current beds record animal activity contemporaneous with nearly continuous sedimentation. Ichnologic studies may also help to distinguish between marine and lacustrine turbidites. Deep-marine turbidites host the Nereites ichnofacies that consists of high diversity of ornate grazing traces and graphoglyptids, recording highly specialized feeding strategies developed to solve the problem of the scarcity of food in the deep sea. Deep lacustrine environments contain the Mermia ichnofacies, which is dominated by unspecialized grazing and feeding traces probably related to the abundance and accessibility of food in lacustrine systems. The lower diversity of lacustrine ichnofaunas in comparison with deep-sea assemblages more likely reflects lower species diversity as a consequence of less stable conditions. Increase of depth and extent of bioturbation through geologic time produced a clear signature in the ichnofabric record of lacustrine facies. Paleozoic lacustrine ichnofaunas are typically dominated by surface trails with little associated bioturbation. During the Mesozoic, bioturbation depth was higher in lake margin facies than in fully lacustrine deposits. While significant degrees of bioturbation were attained in lake margin facies during the Triassic, major biogenic disruption of primary bedding in subaqueous lacustrine deposits did not occur until the Cretaceous.     

Infaunal communities and tiering in Early Palaeozoic nearshore clastic environments: trace-fossil evidence from the Cambro-Ordovician of New South Wales

Understanding of the palaeoecology of Early Palaeozoic shallow-marine communities in sandy habitats is incomplete. Reasons for this include the poor preservation of body fossils and the dominance of Skolithos piperock in sandstones of that age. Cambrian and Ordovician deposits preserving diverse nearshore trace-fossil assemblages are therefore of critical importance for assessing the early evolution of marine ecosystems. The basal part of the Cambro-Ordovician Bynguano Formation of the Mootwingee area (New South Wales, Australia) is a series of interbedded sandstones and mudstones, which were deposited under nearshore conditions. These strata provide an early example of the Arenicolites ichnofacies. Two ichnocoenoses are distinguished in the sandstones of this unit. A 'predepositional' ichnocoenosis, which reflects the benthic community prior to episodes of sand deposition, includes dense aggregations of Rusophycus with rare Planolites. The 'postdepositional' ichnocoenosis is more diverse and includes Thalassinoides, Arenicolites (various types), Monocraterion, Skolithos, Trichichnus , and epichnial grooves. The tiered structure developed in this ichnocoenosis is preserved as a 'frozen tiered profile' and is characterised by a Thalassinoides tier, 10–30 cm in depth, which is cross-cut by Skolithos and Arenicolitesin the middle tier, and by Arenicolites and Trichichnusin the shallowest tier. The pattern of tiering indicates that a complex ecosystem of opportunistic organisms, capable of exploiting shifting substrates, had evolved by the earliest Ordovician. □ Predepositional, postdepositional frozen tier profile, ichnocoenosis, nearshore clastics, RUSOPHYCUS, Cambro-Ordovician, Bynguano Formation, Mootwingee, New South Wales.