Jurassic paleosurfaces with fecal mounds reveal the last supper of arenicolid worms.

Exceptional paleosurfaces preserving fecal casting mounds occur in the Upper Jurassic Lastres Formation of Spain. As in modern shorelines, these biogenic structures are associated with straight to sinuous-crested ripples showing the interplay of biological and physical processes in a low-energy marine environment. These trace fossils display characteristics, distribution, and densities like those of modern arenicolid populations (approximately 35 specimens per m2). Under close examination, these fecal casting mounds are morphologically undistinguishable from those produced by recent arenicolids (e.g. Arenicola marina, Abarenicola pacifica), providing evidence of the presence of these polychaetes in the Late Jurassic. As their modern counterparts, fossil arenicolids very likely modified their environment generating a seabed topography and impacting ancient benthic communities, sediment characteristics, and sediment biogeochemistry. Although the presence of oxic microhabitats and biogeochemical processes cannot be accurately measured in the fossil record, comparison with the work of modern populations allows to make inferences on sediment reworking and bioirrigation potential. In addition, association with grazing trails supports the idea of fertilization and modulation of food resources to other species. These paleosurfaces underscore the significance of high-fidelity snapshots in the fossil record (true substrates) to reconstruct past ecologies and sediment biogeochemistry. A new ichnotaxon, Cumulusichnus asturiensis n. igen. and n. isp., is defined.

Increased habitat segregation at the dawn of the Phanerozoic revealed by correspondence analysis of bioturbation.

The Agronomic Revolution of the early Cambrian refers to the most significant re-structuration of the benthic marine ecosystem in life history. Using a global compilation of trace-fossil records across the Ediacaran-Cambrian transition, this paper investigates the relationship between the benthos and depositional environments prior to, during, and after the Agronomic Revolution to shed light on habitat segregation via correspondence analysis. The results of this analysis characterize Ediacaran mobile benthic bilaterians as facies-crossing and opportunistic, with low levels of habitat specialization. In contrast, the Terreneuvian and Cambrian Series 2 reveal progressive habitat segregation, parallel to matground environmental restriction. This event was conducive to the establishment of distinct endobenthic communities along the marine depositional profile, showing that the increase in styles of animal-substrate interactions was expressed by both alpha and beta ichnodiversity. Habitat segregation at the dawn of the Phanerozoic may illustrate an early extension of the trophic group amensalism at community scale.

Deep-sea Ordovician lingulide brachiopods and their associated burrows suggest an early colonization of proximal turbidite systems.

Trace fossils from Ordovician deep-marine environments are typically produced by a shallow endobenthos adapted to live under conditions of food scarcity by means of specialized grazing, farming, and trapping strategies, preserved in low-energy intermediate to distal zones of turbidite systems. High-energy proximal zones have been considered essentially barren in the early Paleozoic. We report here the first trace and body fossils of lingulide brachiopods in deep-marine environments from an Upper Ordovician turbidite channel-overbank complex in Asturias, Spain. Body and trace fossils are directly associated, supporting the interpretation of a lingulide tracemaker. Ellipsoidal cross-section, cone-in-cone spreite, and spade morphologies suggest the specimens belong to Lingulichnus verticalis. The oblique orientation in both trace and body fossils is the result of tectonic deformation. The organisms were suspension feeders showing escape, dwelling, and equilibrium behaviours controlled by sedimentation rates associated with turbidite deposition. These trace fossils and their in situ producers represent the oldest evidence of widespread endobenthos colonization in high-energy, proximal areas of turbidite systems, expanding the bathymetric range of Lingulichnus and the variety of behaviours and feeding styles in early Paleozoic deep-marine environments.

Probability-based preservational variations within the early Cambrian Chengjiang biota (China).

The Chengjiang biota (Yunnan Province, China) is a treasure trove of soft-bodied animal fossils from the earliest stages of the Cambrian explosion. The mechanisms contributing to its unique preservation, known as the Burgess Shale-type preservation, are well understood. However, little is known about the preservation differences between various animal groups within this biota. This study compares tissue-occurrence data of 11 major animal groups in the Chengjiang biota using a probabilistic methodology. The fossil-based data from this study is compared to previous decay experiments. This shows that all groups are not equally preserved with some higher taxa more likely to preserve soft tissues than others. These differences in fossil preservation between taxa can be explained by the interaction of biological and environmental characteristics. A bias also results from differential taxonomic recognition, as some taxa are easily recognized from even poorly preserved fragments while other specimens are difficult to assign to higher taxa even with exquisite preservation.

Animal-substrate interactions preserved in ancient lagoonal chalk.

Trace-fossil assemblages reflect the response of the benthos to sets of paleoenvironmental conditions during and immediately after sedimentation. Trace fossils have been widely studied in pelagic shelf and deep-sea chalk deposits from around the globe but never documented from ancient lagoonal chalk successions. Here we report the first detailed ichnologic analysis of a lagoonal chalk unit, using as an example the Upper Cretaceous Buda Formation from the Texas Gulf Coast Basin. In this unit, variable interconnection with the open ocean, accompanied by marked fluctuations in physicochemical parameters inherent to lagoonal circulation (e.g., salinity, hydrodynamic energy, bottom-water oxygenation), highly influenced the resultant trace-fossil content of the chalk. These lagoonal chalk deposits contain twenty ichnotaxa, displaying a clear dominance of Thalassinoides isp. and Chondrites isp., which are present in most of the bioturbated strata. The dominance of Thalassinoides isp., both in softgrounds as an element of the Cruziana Ichnofacies and in firmgrounds as a component of the Glossifungites Ichnofacies, highlights similarities with trace-fossil assemblages from shallow-water shelf-sea chalks. In contrast to both (open) shallow-water shelf-sea chalks and deep-sea chalks, the Buda Formation chalk exhibits more diverse assemblages and sharp fluctuations in ichnodiversity and ichnodisparity during relatively short periods of time. The increased ichnodiversity and ichnodisparity in this lagoonal chalk (in comparison with its open ocean counterparts) may reflect a complex interplay of taphonomic (i.e., incomplete bioturbation allowing preservation of shallow-tier trace fossils and ecologic (i.e., increased spatial environmental heterogeneity in the carbonate lagoonal setting) factors.

Assessing the expansion of the Cambrian Agronomic Revolution into fan-delta environments.

The intensity, extent, and ecosystem-level impact of bioturbation (i.e. Agronomic Revolution) at the dawn of the Phanerozoic is a hotly debated issue. Middle Cambrian fan-delta deposits in southwestern Saskatchewan provide insights into the paleoenvironmental extent of the Agronomic Revolution into marginal-marine environments. The studied deposits reveal that several environmental stressors had direct impact on trace-fossil distribution and bioturbation intensities in Cambrian fan deltas. Basal and proximal subaerial deposits are characterized by very coarse grain size and absence of bioturbation. Mid-fan and fan-toe deposits were formed under subaqueous conditions and are characterized by rapid bioturbation events in between sedimentation episodes when environmental stressors were ameliorated, providing evidence of a significant landward expansion of the Agronomic Revolution. Transgressive marine deposits accumulated after the abandonment of the fan-delta system display high levels of bioturbation intensity, reflecting stable environmental conditions that favored endobenthic colonization. The presence of intense bioturbation in both subaqueous fan delta and transgressive deposits provides further support to the view that Cambrian levels of biogenic mixing were high, provided that stable environmental conditions were reached. Our study underscores the importance of evaluating sedimentary facies changes to assess the impact of environmental factors prior to making evolutionary inferences.

The Invasion of the Land in Deep Time: Integrating Paleozoic Records of Paleobiology, Ichnology, Sedimentology, and Geomorphology.

The invasion of the land was a complex, protracted process, punctuated by mass extinctions, that involved multiple routes from marine environments. We integrate paleobiology, ichnology, sedimentology, and geomorphology to reconstruct Paleozoic terrestrialization. Cambrian landscapes were dominated by laterally mobile rivers with unstable banks in the absence of significant vegetation. Temporary incursions by arthropods and worm-like organisms into coastal environments apparently did not result in establishment of continental communities. Contemporaneous lacustrine faunas may have been inhibited by limited nutrient delivery and high sediment loads. The Ordovician appearance of early land plants triggered a shift in the primary locus of the global clay mineral factory, increasing the amount of mudrock on the continents. The Silurian-Devonian rise of vascular land plants, including the first forests and extensive root systems, was instrumental in further retaining fine sediment on alluvial plains. These innovations led to increased architectural complexity of braided and meandering rivers. Landscape changes were synchronous with establishment of freshwater and terrestrial arthropod faunas in overbank areas, abandoned fluvial channels, lake margins, ephemeral lakes, and inland deserts. Silurian-Devonian lakes experienced improved nutrient availability, due to increased phosphate weathering and terrestrial humic matter. All these changes favoured frequent invasions to permament establishment of jawless and jawed fishes in freshwater habitats and the subsequent tetrapod colonization of the land. The Carboniferous saw rapid diversification of tetrapods, mostly linked to aquatic reproduction, and land plants, including gymnosperms. Deeper root systems promoted further riverbank stabilization, contributing to the rise of anabranching rivers and braided systems with vegetated islands. New lineages of aquatic insects developed and expanded novel feeding modes, including herbivory. Late Paleozoic soils commonly contain pervasive root and millipede traces. Lacustrine animal communities diversified, accompanied by increased food-web complexity and improved food delivery which may have favored permanent colonization of offshore and deep-water lake environments. These trends continued in the Permian, but progressive aridification favored formation of hypersaline lakes, which were stressful for colonization. The Capitanian and end-Permian extinctions affected lacustrine and fluvial biotas, particularly the invertebrate infauna, although burrowing may have allowed some tetrapods to survive associated global warming and increased aridification.

The Chengjiang Biota inhabited a deltaic environment.

The Chengjiang Biota is the earliest Phanerozoic soft-bodied fossil assemblage offering the most complete snapshot of Earth's initial diversification, the Cambrian Explosion. Although palaeobiologic aspects of this biota are well understood, the precise sedimentary environment inhabited by this biota remains debated. Herein, we examine a non-weathered core from the Yu'anshan Formation including the interval preserving the Chengjiang Biota. Our data indicate that the succession was deposited as part of a delta influenced by storm floods (i.e., produced by upstream river floods resulting from ocean storms). Most Chengjiang animals lived in an oxygen and nutrient-rich delta front environment in which unstable salinity and high sedimentation rates were the main stressors. This unexpected finding allows for sophisticated ecological comparisons with other Burgess Shale-type deposits and emphasizes that the long-held view of Burgess Shale-type faunas as snapshots of stable distal shelf and slope communities needs to be revised based on recent sedimentologic advances.

Contrasting Early Ordovician assembly patterns highlight the complex initial stages of the Ordovician Radiation.

The Early Ordovician is a key interval for our understanding of the evolution of life on Earth as it lays at the transition between the Cambrian Explosion and the Ordovician Radiation and because the fossil record of the late Cambrian is scarce. In this study, assembly processes of Early Ordovician trilobite and echinoderm communities from the Central Anti-Atlas (Morocco), the Montagne Noire (France), and the Cordillera Oriental (Argentina) are explored. The results show that dispersal increased diachronically in trilobite communities during the Early Ordovician. Dispersal did not increase for echinoderms. Dispersal was most probably proximally triggered by the planktic revolution, the fall in seawater temperatures, changes in oceanic circulation, with an overall control by tectonic frameworks and phylogenetic constraints. The diachronous increase in dispersal within trilobite communities in the Early Ordovician highlights the complexity of ecosystem structuring during the early stages of the Ordovician Radiation. As Early Ordovician regional dispersal was followed by well-documented continental dispersal in the Middle/Late Ordovician, it is possible to consider that alongside a global increase in taxonomic richness, the Ordovician Radiation is also characterized by a gradual increase in dispersal.

Were all trilobites fully marine? Trilobite expansion into brackish water during the early Palaeozoic.

Trilobites, key components of early Palaeozoic communities, are considered to have been invariably fully marine. Through the integration of ichnological, palaeobiological, and sedimentological datasets within a sequence-stratigraphical framework, we challenge this assumption. Here, we report uncontroversial trace and body fossil evidence of their presence in brackish-water settings. Our approach allows tracking of some trilobite groups foraying into tide-dominated estuaries. These trilobites were tolerant to salinity stress and able to make use of the ecological advantages offered by marginal-marine environments migrating up-estuary, following salt wedges either reflecting amphidromy or as euryhaline marine wanderers. Our data indicate two attempts of landward exploration via brackish water: phase 1 in which the outer portion of estuaries were colonized by olenids (Furongian-early late Tremadocian) and phase 2 involving exploration of the inner to middle estuarine zones by asaphids (Dapingian-Darriwilian). This study indicates that tolerance to salinity stress arose independently among different trilobite groups.

Quantifying ecospace utilization and ecosystem engineering during the early Phanerozoic-The role of bioturbation and bioerosion.

The Cambrian explosion (CE) and the great Ordovician biodiversification event (GOBE) are the two most important radiations in Paleozoic oceans. We quantify the role of bioturbation and bioerosion in ecospace utilization and ecosystem engineering using information from 1367 stratigraphic units. An increase in all diversity metrics is demonstrated for the Ediacaran-Cambrian transition, followed by a decrease in most values during the middle to late Cambrian, and by a more modest increase during the Ordovician. A marked increase in ichnodiversity and ichnodisparity of bioturbation is shown during the CE and of bioerosion during the GOBE. Innovations took place first in offshore settings and later expanded into marginal-marine, nearshore, deep-water, and carbonate environments. This study highlights the importance of the CE, despite its Ediacaran roots. Differences in infaunalization in offshore and shelf paleoenvironments favor the hypothesis of early Cambrian wedge-shaped oxygen minimum zones instead of a horizontally stratified ocean.

The rise and early evolution of animals: where do we stand from a trace-fossil perspective?

The trace-fossil record provides a wealth of information to track the rise and early evolution of animals. It comprises the activity of both hard- and soft-bodied organisms, is continuous through the Ediacaran (635-539 Ma)- Cambrian (539-485 Ma) transition, yields insights into animal behaviour and their role as ecosystem engineers, and allows for a more refined characterization of palaeoenvironmental context. In order to unravel macroevolutionary signals from the trace-fossil record, a variety of approaches is available, including not only estimation of degree of bioturbation, but also analysis of ichnodiversity and ichnodisparity trajectories, and evaluation of the occupation of infaunal ecospace and styles of ecosystem engineering. Analysis of the trace-fossil record demonstrates the presence of motile benthic bilaterians in the Ediacaran, mostly feeding from biofilms. Although Ediacaran trace fossils are simple and emplaced at or immediately below the sediment surface, an increase in ichnofossil complexity, predation pressure, sediment disturbance and penetration depth is apparent during the terminal Ediacaran. Regardless of this increase, a dramatic rise in trace fossil diversity and disparity took place during the earliest Cambrian, underscoring that the novelty of the Fortunian (539-529 Ma) cannot be underestimated. The Fortunian still shows the persistence of an Ediacaran-style matground ecology, but is fundamentally characterized by the appearance of new trace-fossil architectural plans reflecting novel ways of interacting with the substrate. The appearance of Phanerozoic-style benthic ecosystems attests to an increased length and connectivity of the food web and improved efficiency in organic carbon transfer and nutrient recycling. A profound reorganization of the infaunal ecospace is recorded in both high-energy sand-dominated nearshore areas and low-energy mud-dominated offshore environments, during the early Cambrian, starting approximately during Cambrian Age 2 (529-521 Ma), but continuing during the rest of the early Cambrian. A model comprising four evolutionary phases is proposed to synthetize information from the Ediacaran-Cambrian trace-fossil record. The use of a rich ichnological toolbox; critical, systematic and comprehensive evaluation of the Ediacaran-Cambrian trace-fossil record; and high-resolution integration of the ichnological dataset and sedimentological information show that the advent of biogenic mixing was an important factor in fully marine environments at the dawn of the Phanerozoic.

A new insect trackway from the Upper Jurassic-Lower Cretaceous eolian sandstones of São Paulo State, Brazil: implications for reconstructing desert paleoecology.

The new ichnospecies Paleohelcura araraquarensis isp. nov. is described from the Upper Jurassic-Lower Cretaceous Botucatu Formation of Brazil. This formation records a gigantic eolian sand sea (erg), formed under an arid climate in the south-central part of Gondwana. This trackway is composed of two track rows, whose internal width is less than one-quarter of the external width, with alternating to staggered series, consisting of three elliptical tracks that can vary from slightly elongated to tapered or circular. The trackways were found in yellowish/reddish sandstone in a quarry in the Araraquara municipality, São Paulo State. Comparisons with neoichnological studies and morphological inferences indicate that the producer of Paleohelcura araraquarensis isp. nov. was most likely a pterygote insect, and so could have fulfilled one of the ecological roles that different species of this group are capable of performing in dune deserts. The producer could have had a herbivorous or carnivorous diet or been part of the fauna of omnivores, being able to adopt herbivorous, carnivorous, and saprophagous diets when necessary. In modern dune deserts, some species of pterygote insects are detritivores (like Tenebrionidae), relying on organic matter that accumulated among the sand grains of the dunes during dry periods with no plant growth. The presence of additional burrows suggests that the Botucatu paleodesert would have had a detritivorous fauna like this. Based on the interpretation of the ichnofossil producers, it was possible to reconstruct the food web of this paleodesert. All the omnivorous and herbivorous invertebrates and the herbivorous ornithopod dinosaurs made up the primary consumers. These animals were, in turn, the food source for bigger carnivorous or omnivorous animals unable to feed on detritus, like arachnids, possible predatory insects, mammaliaforms, and theropod dinosaurs. The highest trophic level was occupied by larger theropod dinosaurs and mammaliaforms, which, because of their size, could prey upon a wide range of animals. The producer of Paleohelcura araraquarensis isp. nov. could have been a primary consumer if it were an omnivorous detritivore or a herbivore, or a secondary consumer if it were produced by a predatory insect or an omnivore relying on animal biomass. The description of this new trackway expands the knowledge on the faunal composition of the Botucatu paleodesert and provides insights into the ecological relationships in ancient deserts. The presence of these arthropod trackways in Mesozoic eolian deposits helps to trace a continuity between Paleozoic and post-Paleozoic desert ichnofaunas, further reinforcing a single Octopodichnus-Entradichnus Ichnofacies for eolian deposits.

Trace fossil evidence for infaunal moulting in a Middle Devonian non-trilobite euarthropod.

Trace fossils represent the primary source of information on the evolution of animal behaviour through deep time, and provide exceptional insights into complex life strategies that would be otherwise impossible to infer from the study of body parts alone. Here, we describe unusual trace fossils found in marginal-marine, storm- and river-flood deposits from the Middle Devonian Naranco Formation of Asturias (northern Spain) that constitute the first evidence for infaunal moulting in a non-trilobite euarthropod. The trace fossils are preserved in convex hyporelief, and include two main morphological variants that reflect a behavioural continuum. Morphotype 1 consists of a structure that superficially resembles a Rusophycus with an oval outline that possesses a distinctly three lobed axis with an elevated central ridge and regularly spaced transverse furrows that convey the appearance of discrete body segments. The anterior part is the most irregular region of the structure, and it is not always recorded. Morphotype 2 displays more elongated, tubular morphology. Careful observation, however, reveals that it comprises up to three successive morphotype 1 specimens organised in a linear fashion and partially truncating each other. Trilobate morphology and effaced transverse furrows are locally evident, but the predominant morphological feature is the continuous, elevated ridge. The detailed morphology of morphotype 1 and well-preserved, discrete segments of morphotype 2 closely resemble the dorsal exoskeleton of the enigmatic late Carboniferous euarthropod Camptophyllia, suggesting the possible affinities of the producer. Comparisons with patterns of Devonian phacopid trilobite exuviation suggest that the Naranco Formation trace fossils may have been produced by the infaunal activities of an euarthropod that anchored its dorsal exoskeleton in the firm sediment during the body inversion moult procedure. Our findings expand the phylogenetic and environmental occurrence of infaunal moulting in Palaeozoic euarthropods, and suggest a defensive strategy against predation, previously only known from trilobites preserved in open-marine deposits.

The search for an elusive worm in the tropics, the past as a key to the present, and reverse uniformitarianism.

The distribution of trace-making organisms in coastal settings is largely controlled by changes in physicochemical parameters, which in turn are a response to different climatic and oceanographic conditions. The trace fossil Macaronichnus and its modern producers are typical of high-energy, siliciclastic foreshore sands in intermediate- to high-latitude settings characterized by cold-water conditions. However, it has been found in Miocene Caribbean deposits of Venezuela, prompting the hypothesis that upwelling of cold, nutrient-rich waters rather than latitude was the main control of its distribution. To test this hypothesis that was solely based on the fossil record, several trenches and sediment peels were made in two high-energy sand beaches having different oceanographic conditions along the Pacific and Caribbean coasts of the Central American Isthmus. As predicted, the burrows were found only in the highly productive waters of the Pacific coast of Costa Rica in connection with upwelling, while they were absent from the warm, oligotrophic waters of the Caribbean coast of Panama. This finding demonstrates that sometimes the past may be a key to the present, providing one of the few documented examples of reverse uniformitarianism.

Trace fossils associated with Burgess Shale non-biomineralized carapaces: bringing taphonomic and ecological controls into focus.

The association of trace fossils and non-biomineralized carapaces has been reported from Cambrian Lagerstätten worldwide, but the abundance, ichnodiversity, taphonomy and ecological significance of such associations have yet to be fully investigated. Two main end-member hypotheses are explored based on the study of a relatively wide variety of trace fossils preserved associated to Tuzoia carapaces from the middle Cambrian Burgess Shale in British Columbia. In the ecological Tuzoia garden hypothesis, the bacterially enriched surface of carapaces provides opportunities for intricate ecologic interactions among trophic levels. In the taphonomic shielding hypothesis, the trace fossil-carapace association results from preferential preservation of traces as controlled by compaction independent of any association in life. In an attempt to better understand the role of the carapace as a medium for preservation of trace fossils and to evaluate the effects of mechanical stress related to burial, a numerical model was developed. Results indicate that the carapace can shield underlying sediment from mechanical stress for a finite time, differentially protecting trace fossils during the initial phase of burial and compaction. However, this taphonomic model alone fails to fully explain relatively high-density assemblages displaying a diversity of structures spatially confined within the perimeter of carapaces or branching patterns recording re-visitation.

Ediacaran biozones identified with network analysis provide evidence for pulsed extinctions of early complex life.

Rocks of Ediacaran age (~635-541 Ma) contain the oldest fossils of large, complex organisms and their behaviors. These fossils document developmental and ecological innovations, and suggest that extinctions helped to shape the trajectory of early animal evolution. Conventional methods divide Ediacaran macrofossil localities into taxonomically distinct clusters, which may represent evolutionary, environmental, or preservational variation. Here, we investigate these possibilities with network analysis of body and trace fossil occurrences. By partitioning multipartite networks of taxa, paleoenvironments, and geologic formations into community units, we distinguish between biostratigraphic zones and paleoenvironmentally restricted biotopes, and provide empirically robust and statistically significant evidence for a global, cosmopolitan assemblage unique to terminal Ediacaran strata. The assemblage is taxonomically depauperate but includes fossils of recognizable eumetazoans, which lived between two episodes of biotic turnover. These turnover events were the first major extinctions of complex life and paved the way for the Cambrian radiation of animals.

Organism motility in an oxygenated shallow-marine environment 2.1 billion years ago.

Evidence for macroscopic life in the Paleoproterozoic Era comes from 1.8 billion-year-old (Ga) compression fossils [Han TM, Runnegar B (1992) Science 257:232-235; Knoll et al. (2006) Philos Trans R Soc Lond B 361:1023-1038], Stirling biota [Bengtson S et al. (2007) Paleobiology 33:351-381], and large colonial organisms exhibiting signs of coordinated growth from the 2.1-Ga Francevillian series, Gabon. Here we report on pyritized string-shaped structures from the Francevillian Basin. Combined microscopic, microtomographic, geochemical, and sedimentologic analyses provide evidence for biogenicity, and syngenicity and suggest that the structures underwent fossilization during early diagenesis close to the sediment-water interface. The string-shaped structures are up to 6 mm across and extend up to 170 mm through the strata. Morphological and 3D tomographic reconstructions suggest that the producer may have been a multicellular or syncytial organism able to migrate laterally and vertically to reach food resources. A possible modern analog is the aggregation of amoeboid cells into a migratory slug phase in cellular slime molds at times of starvation. This unique ecologic window established in an oxygenated, shallow-marine environment represents an exceptional record of the biosphere following the crucial changes that occurred in the atmosphere and ocean in the aftermath of the great oxidation event (GOE).

Early Cambrian origin of the shelf sediment mixed layer.

The mixed layer of modern oceans is a zone of fully homogenized sediment resulting from bioturbation. The mixed layer is host to complex biogeochemical cycles that directly impact ecosystem functioning, affecting ocean productivity and marine biodiversity. The timing of origin of the mixed layer has been controversial, with estimates ranging from Cambrian to Silurian, hindering our understanding of biogeochemical cycling and ecosystem dynamics in deep time. Here we report evidence from the Global Stratotype Section and Point (GSSP) of the basal Cambrian in the Burin Peninsula of Newfoundland, Canada, showing that a well-developed mixed layer of similar structure to that of modern marine sediments was established in shallow marine settings by the early Cambrian (approximately 529 million years ago). These findings imply that the benthos significantly contributed to establishing new biogeochemical cycles during the Cambrian explosion.

Sediment disturbance by Ediacaran bulldozers and the roots of the Cambrian explosion.

Trace fossils of sediment bulldozers are documented from terminal Ediacaran strata of the Nama Group in Namibia, where they occur in the Spitskop Member of the Urusis Formation (Schwarzrand Subgroup). They consist of unilobate to bilobate horizontal to subhorizontal trace fossils describing scribbles, circles and, more rarely, open spirals and meanders, and displaying an internal structure indicative of active fill. Their presence suggests that exploitation of the shallow infaunal ecospace by relatively large bilaterians was already well underway at the dawn of the Phanerozoic. Efficient burrowing suggests coelom development most likely linked to metazoan body-size increase. These trace fossils are the earliest clear representatives so far recorded of sediment bulldozing, an activity that may have had a negative impact on suspension-feeding and/or osmotroph communities, as well as on matgrounds, representing early examples of ecosystem engineering and trophic-group amensalism. The occurrence of sediment bulldozers may have promoted the establishment of gradients in horizontal and vertical distribution of organic material in connection with spatially heterogeneous environments on the sea floor at a critical time in Earth evolution.