Cambrian Animals Research Articles

The Emu Bay Shale: A unique early Cambrian Lagerstätte from a tectonically active basin.

The Emu Bay Shale (EBS) of South Australia is anomalous among Cambrian Lagerstätten because it captures anatomical information that is rare in Burgess Shale-type fossils, and because of its inferred nearshore setting, the nature of which has remained controversial. Intensive study, combining outcrop and borehole data with a compilation of >25,000 fossil specimens, reveals that the EBS biota inhabited a fan delta complex within a tectonically active basin. Preservation of soft-bodied organisms in this setting is unexpected and further underscores differences between the EBS and other Cambrian Lagerstätten. Environmental conditions, including oxygen fluctuations, slope instability, high suspended sediment concentrations, and episodic high-energy events, inhibited colonization of the lower prodelta by all but a few specialist species but favored downslope transportation and preservation of other largely endemic, shallow-water benthos. The EBS provides extraordinary insight into early Cambrian animal diversity from Gondwana. These results demonstrate how environmental factors determined community composition and provide a framework for understanding this unique Konservat-Lagerstätte.

Spatiotemporal distribution and morphological diversity of the Cambrian Wiwaxia: New insights from South China

Wiwaxia is a well-known Cambrian animal and a relative of annelids and molluscs, characterized by its densely arranged, imbricated dorsal sclerites. Despite its prevalence in the fossil record, the scarcity of articulated specimens leading to persistent uncertainties regarding its morphological diversities and evolutionary history has fueled ongoing investigations. This study presents an additional articulated specimen and isolated sclerites from the Cambrian Stage 4 Douposi Formation in Yunnan Province, South China, representing a new species—Wiwaxia douposiensis sp. nov. The newly described materials, originating from a new Cambrian Konservat-Lagerstätte, exhibit typical Burgess Shale-type preservation. They unveil a distinctive two-order rib pattern and aspect ratios of sclerites. The temporal and spatial distribution of Wiwaxia suggests a potential origination centered in South China during the Cambrian Age 3, with an expanding pattern from shallow to deep waters and from warm-humid to dry-cold climates during the Cambrian Age 4 to Drumian. Statistical and phylogenetic analyses highlight the dual aspects of conservative aspect ratios in sclerites, and morphological variations observed at the species levels within Wiwaxia.

Rapid volcanic ash entombment reveals the 3D anatomy of Cambrian trilobites.

Knowledge of Cambrian animal anatomy is limited by preservational processes that result in compaction, size bias, and incompleteness. We documented pristine three-dimensional (3D) anatomy of trilobites fossilized through rapid ash burial from a pyroclastic flow entering a shallow marine environment. Cambrian ellipsocephaloid trilobites from Morocco are articulated and undistorted, revealing exquisite details of the appendages and digestive system. Previously unknown anatomy includes a soft-tissue labrum attached to the hypostome, a slit-like mouth, and distinctive cephalic feeding appendages. Our findings resolve controversy over whether the trilobite hypostome is the labrum or incorporates it and establish crown-group euarthropod homologies in trilobites. This occurrence of moldic fossils with 3D soft parts highlights volcanic ash deposits in marine settings as an underexplored source for exceptionally preserved organisms.

Evidence for low sulfate and anoxic deep waters in early Cambrian

The role of environmental change as a significant force driving biological evolution during the Ediacaran–Cambrian transition remains a subject of extensive debate. In this study, we present high-resolution C-isotope data of organic matter (δ13Corg) and multiple S-isotope compositions of pyrite (δ34Spy and Δ33Spy) obtained from a continuous drill core (ZK1505) in South China, spanning the interval from the Ediacaran to Cambrian Stage 2. Our δ13Corg and δ34Spy data, combined with published data from four sections deposited across a shelf-to-basin transect, reveal substantial gradient from shallow to deep water in δ13Corg and sustained positive δ34Spy values in the early Cambrian. The vertical δ13Corg gradient suggests chemical stratification of the early Cambrian ocean, characterized by oxic shallow water and anoxic deep water. The sustained positive δ34Spy values in the same period suggest low seawater sulfate concentrations. Furthermore, we observed intermittent occurrences of negative Δ33Spy values coupled with positive δ34Spy, suggesting a dominant deep-water anoxia during early Cambrian in the Yangtze Block. We propose that protracted deep-water anoxia may have contributed to the extinction of the Ediacaran biota and played a significant role in the evolution of Cambrian animals.

Chancelloriids from the Cambrian (Stage 4) Balang Lagerstätte of South China and a reappraisal of their diversification in South China

Chancelloriids are an enigmatic group of Cambrian animals characterized by radially symmetrical bodies equipped with spiny sclerites. Although they were major components of many benthic communities, current understanding on how this group diversified during the Cambrian remains limited. Clarifying these confusions about the diversification of chancelloriids requires new information provided by new studies of previously described and/or new material. Considering the emerging scenario of chancelloriids, the sack-like chancelloriids from the Stage 4 Balang Lagerstätte are restudied on the basis of previous collections, which are herein separated from Chancelloria eros Walcott, 1920, and revised as Archiasterella acuminata nov. sp. and Chancelloriidae gen. et sp. indet. The Ar. acuminata specimen is well preserved and does show its scleritome and sclerite characters, while the Chancelloriidae gen. et sp. indet. specimen is poorly preserved and its scleritome and sclerite characters are difficult to distinguish. By combining information from previous studies and the present research, data on chancelloriid occurrences in South China indicate that chancelloriids had once diversified in the Cambrian South China, evidenced by high-level richness of sclerite-based taxa during the late Fortunian to the middle Stage 3, and of sclerite formulas during the late Stage 2 to the Wuliuan. The result is inconsistent with the recent viewpoint that there were two flourishing ages of chancelloriids in South China. Although more evidence is needed to support our results, the present study offers insights in understanding the evolutionary dynamics of chancelloriids.

Hg isotope evidence for oceanic oxygenation during the Cambrian Explosion

The Cambrian explosion is a critical evolutionary milestone in life history, but the mechanistic relationship between the Cambrian explosion, rising atmospheric and oceanic oxygen levels, and the primary productivity remains controversial. Here, we present new mercury (Hg) isotope data from Cambrian marine sediments of the ∼521–514 Ma Yu’anshan and Shuijingtuo Formations (Nanhua Basin, South China), which preserve the famous ∼518 Ma Chengjiang and Qingjiang Cambrian Biotas, respectively. We find evidence that, prior to Cambrian animal diversification, local terrestrial and/or atmospheric inputs are important drivers of Hg enrichment in the shallow shelf Yu’anshan and Shuijintuo Formations. Elevated Hg and total organic carbon (TOC) concentrations of coeval deeper shelf and slope sections of the Niutitang Formation are mainly attributed to upwelling of Hg associated with dissolved organic carbon (Hg-DOC) from pelagic seawater, unrelated to locally high primary productivity. During the deposition of fossiliferous strata, negative shifts of Δ199Hg and δ202Hg values in the shallow water shelf shales from the Shuijingtuo Formation, combined with similar Hg isotope characteristics in the deeper shelf and slope shales, demonstrate a significant shallow and deep-water oxygenation event took place in the Nanhua Basin. A comparable negative shift of Δ199Hg values occurs in coeval Indian Craton sediments, suggesting this oxygenation event could be regional, or possibly even global, in scale. Our newly-collected Hg isotope data provide strong evidence that rising molecular oxygen levels in both surface and deep seawater are associated with enhanced marine primary productivity and could be a critical driver of the Cambrian explosion. Finally, we also argue that Hg isotopes are an emerging and promising redox proxy for studies of Precambrian seawater, but the chemical response of Hg to atmospheric and ocean oxygenation requires further calibration.

Fluctuation in redox conditions and the evolution of early Cambrian life constrained by nitrogen isotopes in the middle Yangtze Block, South China

AbstractThe Ediacaran–Cambrian (E-C) transition (∼542–517 Ma) witnessed the rapid evolution of Cambrian animals, which was accompanied by carbon cycling anomalies and a significant increase in the concentration of oxygen in Earth’s atmosphere. The mechanisms stimulating the evolution of complex eukaryotes, however, remain problematic, especially concerning the link between biological evolution and contemporaneous changes in the oceanic environment. In this study, integrated δ13Ccarb–δ13Corg–δ15N compositions were analysed from the YD-4 core samples to understand redox fluctuations and nitrogen cycling of the middle Yangtze Block across the E-C transition. Two negative δ13Ccarb excursions (N1 and N2) and a positive δ13Ccarb excursion (P1) are identified from the studied samples and are supposedly of primary origin. Constrained by of the U-Pb age, biolithology and pattern of isotopic variation, N1, P1 and N2 are comparable to the Basal Cambrian Carbon Isotope Excursion (BACE), Zhujiaqing Carbon Isotope Excursion (ZHUCE) and Shiyantou Carbon Isotope Excursion (SHICE). We interpreted the decreased δ15N values in this study as resulting from intensified atmospheric nitrogen fixation driven by enhanced denitrification associated with expanded marine anoxia, as well as partial ammonium assimilation, while increased δ15N values suggest weakened denitrification associated with an amplified oxic water mass. The temporal coincidence of N1 and N2, with two episodes of negative δ15N excursions, and of P1, with a positive δ15N excursion, suggests that variable oceanic redox conditions and nitrogen bioavailability may have influenced the evolution of the Cambrian eukaryote-dominated community.

Large euarthropod carapaces from a high latitude Cambrian (Drumian) deposit in Spain.

Deposits preserving non-biomineralized tissues and animals provide an unrivalled opportunity to study the evolution and radiation of early animal life. Numerous sites of Cambrian age are known from North America (Laurentia) and South China (East Gondwana), which provide a high resolution picture of the fauna at low latitudes. By contrast, our knowledge of Cambrian animals from higher latitudes is relatively poor. This patchiness in our knowledge of animal life during the radiation of animals in the Cambrian period limits our ability to understand and detect palaeogeographic trends and does not provide a full appreciation of animal diversity at this time. Here we report a new middle Cambrian (Drumian) site preserving lightly sclerotized euarthropod carapaces, sponges and palaeoscolecids near the village of Mesones de Isuela in the Iberian Chains (Spain). We describe three bivalved euarthropod carapace morphs, two comparable to those described from the only other high latitude Drumian deposit, the Jince Formation (Czechia), and one distinct from previous discoveries. These new findings highlight the importance of high latitude Gondwana Konservat Lagerstatten for understanding the palaeogeographical aspect of the radiation of early animals and suggest that bivalved euarthropods at high latitudes were larger than those at lower latitudes during the Cambrian.

Raptorial appendages of the Cambrian apex predator Anomalocaris canadensis are built for soft prey and speed.

The stem-group euarthropod Anomalocaris canadensis is one of the largest Cambrian animals and is often considered the quintessential apex predator of its time. This radiodont is commonly interpreted as a demersal hunter, responsible for inflicting injuries seen in benthic trilobites. However, controversy surrounds the ability of A. canadensis to use its spinose frontal appendages to masticate or even manipulate biomineralized prey. Here, we apply a new integrative computational approach, combining three-dimensional digital modelling, kinematics, finite-element analysis (FEA) and computational fluid dynamics (CFD) to rigorously analyse an A. canadensis feeding appendage and test its morphofunctional limits. These models corroborate a raptorial function, but expose inconsistencies with a capacity for durophagy. In particular, FEA results show that certain parts of the appendage would have experienced high degrees of plastic deformation, especially at the endites, the points of impact with prey. The CFD results demonstrate that outstretched appendages produced low drag and hence represented the optimal orientation for speed, permitting acceleration bursts to capture prey. These data, when combined with evidence regarding the functional morphology of its oral cone, eyes, body flaps and tail fan, suggest that A. canadensis was an agile nektonic predator that fed on soft-bodied animals swimming in a well-lit water column above the benthos. The lifestyle of A. canadensis and that of other radiodonts, including plausible durophages, suggests that niche partitioning across this clade influenced the dynamics of Cambrian food webs, impacting on a diverse array of organisms at different sizes, tiers and trophic levels.

Ultrastructure reveals ancestral vertebrate pharyngeal skeleton in yunnanozoans.

Pharyngeal arches are a key innovation that likely contributed to the evolution of the jaws and braincase of vertebrates. It has long been hypothesized that the pharyngeal (branchial) arch evolved from an unjointed cartilaginous rod in vertebrate ancestors such as that in the nonvertebrate chordate amphioxus, but whether such ancestral anatomy existed remains unknown. The pharyngeal skeleton of controversial Cambrian animals called yunnanozoans may contain the oldest fossil evidence constraining the early evolution of the arches, yet its correlation with that of vertebrates is still disputed. By examining additional specimens in previously unexplored techniques (for example, x-ray microtomography, scanning and transmission electron microscopy, and energy dispersive spectrometry element mapping), we found evidence that yunnanozoan branchial arches consist of cellular cartilage with an extracellular matrix dominated by microfibrils, a feature hitherto considered specific to vertebrates. Our phylogenetic analysis provides further support that yunnanozoans are stem vertebrates.

A three-eyed radiodont with fossilized neuroanatomy informs the origin of the arthropod head and segmentation

In addition to being among the most iconic and bizarre-looking Cambrian animals, radiodonts are a groupthat offers key insight into the acquisition of the arthropod body plan by virtue of their phylogenetic divergence prior to all living members of the phylum. Nonetheless, radiodont fossils are rare and often fragmentary, and contentions over their interpretation have hindered resolution of important evolutionary conundrums. Here, we describe 268 specimens of Stanleycaris hirpex from the Cambrian Burgess Shale, including many exceptionally preserved whole-body specimens, informing the most complete reconstruction of a radiodont to date. The trunk region of Stanleycaris has up to 17 segments plus two pairs of filiform caudal blades. The recognition of dorsal sclerotic segmentation of the trunk cuticle and putative unganglionated nerve cords provides new insight into the relative timing of acquisition of segmental traits, the epitome of the arthropod body plan. In addition to the pair of stalked lateral eyes, the short head unexpectedly bears a large median eye situated behind a preocular sclerite on an anteriorly projecting head lobe. Upon re-evaluation, similar median eyes can be identified in other Cambrian panarthropods demonstrating a deep evolutionary continuity. The exquisitely preserved brain of Stanleycaris is consistent with the hypothesized deutocerebral innervation of the frontal appendages, reconciling neuroanatomical evidence with external morphology in support of an ancestrally bipartite head and brain for arthropods. We propose that the integration of this bipartite head prior to the acquisition of most segmental characters exclusively in the arthropod trunk may help explain its developmental differentiation.

A new approach for investigating spatial relationships of ichnofossils: a case study of Ediacaran–Cambrian animal traces

AbstractTrace fossils record foraging behaviors, the search for resources in patchy environments, of animals in the rock record. Quantification of the strength, density, and nature of foraging behaviors enables the investigation of how these may have changed through time. Here, we present a novel approach to explore such patterns using spatial point process analyses to quantify the scale and strength of ichnofossil spatial distributions on horizontal bedding planes. To demonstrate the utility of this approach, we use two samples from the terminal Ediacaran Shibantan Member in South China (between 551 and 543 Ma) and the early Cambrian Nagaur Sandstone in northwestern India (between 539 and 509 Ma). We find that ichnotaxa on both surfaces exhibited significant nonhomogeneous lateral patterns, with distinct levels of heterogeneity exhibited by different types of trace fossils. In the Shibantan, two ichnotaxa show evidence for mutual positive aggregation over a shared resource, suggesting the ability to focus on optimal resource areas. Trace fossils from the Nagaur Sandstone exhibit more sophisticated foraging behavior, with greater niche differentiation. Critically, mark correlation functions highlight significant spatial autocorrelation of trace fossil orientations, demonstrating the greater ability of these Cambrian tracemakers to focus on optimal patches. Despite potential limitations, these analyses hint at changes in the development and optimization of foraging at the Ediacaran/Cambrian transition and highlight the potential of spatial point process analysis to tease apart subtle differences in behavior in the trace fossil record.

Global Hg cycle over Ediacaran–Cambrian transition and its implications for environmental and biological evolution

The Ediacaran–Cambrian (E–C) transition witnessed remarkable environmental changes, the extinction of the Ediacaran biota, and subsequent rapid diversification of Cambrian animals. However, the linkages between environmental and biological evolution are still under debate at this critical time. Here, we present new Hg abundance and Hg isotopes in sediments from South China, which are then combined with those published from the Indian craton to explore the co-evolution of environment and complex life during the E–C transition. In both areas, high Hg/TOC ratios and near-zero Δ199Hg of the Late Ediacaran sediments suggest enhanced volcanism, whereas relatively high Hg/TOC ratios and positive Δ199Hg shift upsection indicate volcanic-sourced atmospheric Hg(II) deposition in the earliest Cambrian. The dramatically decreasing Hg/TOC ratios and positive Δ199Hg of early Cambrian Age 2 to 3 sediments indicate scavenging of dissolved seawater Hg(II) by organic matter particulates. Our Hg results suggest volcanism may have likely played a significant role in the extinction of Ediacaran biota and global negative carbon excursions near the E–C boundary. Furthermore, our Hg data provides new evidence of extensive OM burial in the early Cambrian Age 2 to 3 oceans, leading to a rapid increase of Earth-surface O2 levels that coincided with appearance of more complex large-body animals. Our study provides new insights of the global Hg cycle into the co-evolution of the environment and complex life at this critical time.

Tube-dwelling in early animals exemplified by Cambrian scalidophoran worms

BackgroundThe radiation of ecdysozoans (moulting animals) during the Cambrian gave rise to panarthropods and various groups of worms including scalidophorans, which played an important role in the elaboration of early marine ecosystems. Although most scalidophorans were infaunal burrowers travelling through soft sediment at the bottom of the sea, Selkirkia lived inside a tube.ResultsWe explore the palaeobiology of these tubicolous worms, and more generally the origin and evolutionary significance of tube-dwelling in early animals, based on exceptionally preserved fossils from the early Cambrian Chengjiang Lagerstätte (Stage 3, China) including a new species, Selkirkia transita sp. nov. We find that the best phylogenetic model resolves Selkirkia as a stem-group priapulid. Selkirkia secreted a protective cuticular thickening, the tube, inside which it was able to move during at least part of its life. Partly based on measured growth patterns, we construe that this tube was separated from the trunk during a moulting process that has no direct equivalent in other scalidophorans. Although the ontogeny of Selkirkia is currently unknown, we hypothesize that its conical tube might have had the same ecological function and possibly even deep development origin as the lorica, a protective cuticular thickening found in larval priapulids and adult loriciferans. Selkirkia is seen as a semi-sedentary animal capable of very shallow incursions below the water/sediment interface, possibly for feeding or during the tube-secreting phase. Brachiopod epibionts previously reported from the Xiaoshiba Lagerstätte (ca. 514 Ma) also presumably occur in Selkirkia sinica from Chengjiang (ca. 518 Ma).ConclusionsOur critical and model-based approach provides a new phylogenetic framework for Scalidophora, upon which to improve in order to study the evolution of morphological characters in this group. Tube-dwelling is likely to have offered Selkirkia better protection and anchoring to sediment and has developed simultaneously in other Cambrian animals such as hemichordates, annelids or panarthropods. Often lost in modern representatives in favour of active infaunal lifestyles, tube-dwelling can be regarded as an early evolutionary response of various metazoans to increasing environmental and biological pressure in Cambrian marine ecosystems.

Vertically migrating Isoxys and the early Cambrian biological pump.

The biological pump is crucial for transporting nutrients fixed by surface-dwelling primary producers to demersal animal communities. Indeed, the establishment of an efficient biological pump was likely a key factor enabling the diversification of animals over 500 Myr ago during the Cambrian explosion. The modern biological pump operates through two main vectors: the passive sinking of aggregates of organic matter, and the active vertical migration of animals. The coevolution of eukaryotes and sinking aggregates is well understood for the Proterozoic and Cambrian; however, little attention has been paid to the establishment of the vertical migration of animals. Here we investigate the morphological variation and hydrodynamic performance of the Cambrian euarthropod Isoxys. We combine elliptical Fourier analysis of carapace shape with computational fluid dynamics simulations to demonstrate that Isoxys species likely occupied a variety of niches in Cambrian oceans, including vertical migrants, providing the first quantitative evidence that some Cambrian animals were adapted for vertical movement in the water column. Vertical migration was one of several early Cambrian metazoan innovations that led to the biological pump taking on a modern-style architecture over 500 Myr ago.

Quantitative comparison of geological data and model simulations constrains early Cambrian geography and climate.

Marine ecosystems with a diverse range of animal groups became established during the early Cambrian (~541 to ~509 Ma). However, Earth’s environmental parameters and palaeogeography in this interval of major macro-evolutionary change remain poorly constrained. Here, we test contrasting hypotheses of continental configuration and climate that have profound implications for interpreting Cambrian environmental proxies. We integrate general circulation models and geological observations to test three variants of the ‘Antarctocentric’ paradigm, with a southern polar continent, and an ‘equatorial’ configuration that lacks polar continents. This quantitative framework can be applied to other deep-time intervals when environmental proxy data are scarce. Our results show that the Antarctocentric palaeogeographic paradigm can reconcile geological data and simulated Cambrian climate. Our analyses indicate a greenhouse climate during the Cambrian animal radiation, with mean annual sea-surface temperatures between ~9 °C to ~19 °C and ~30 °C to ~38 °C for polar and tropical palaeolatitudes, respectively.

Pyritized preservation of chancelloriids from the Cambrian Stage 3 of South China and implications for biomineralization

The enigmatic Cambrian animal chancelloriids were discovered in a wide range of taphonomic settings; however, preservation of biomineralized sclerite microstructure was solely known from secondarily phosphatized skeletal remains. Here, we investigate a uniquely pyritized chancelloriid from the lower Cambrian Guojiaba Formation in southern Shaanxi Province, China, using a combination of advanced analytic techniques. Results of the energy dispersive spectroscopy (EDS), X-Ray Fluorescence (XRF), and Raman spectrum show that the sclerites and scleritomes are preserved as pyritized internal moulds with a calcitic outer layer. The outer layer enveloping the internal moulds likely represents the recrystallized counterpart of the original biomineralized sclerite wall. Distinctive fibrous microstructures are discovered in the sclerites, which echo the features seen in the phosphatized fossils of chancelloriids. The typical microstructure, along with the recrystallized calcite, corroborate the interpretation that chancelloriid sclerites were originally constructed by fibrous aragonite. The stability of the microstructure and mineral composition in both carbonate and siliciclastic backgrounds indicate that chancelloriids were adapted to exploit aragonitic fibres to build their skeletons regardless of the change of their living environments.

Priapulid worms from the Cambrian of China shed light on reproduction in early animals

In the recent years, exceptional fossil sites have revealed astonishing details on the anatomy, lifestyles and behaviour of Cambrian animals but surprisingly, very little is known about one of their most vital features, reproduction. We describe here in situ eggs (clusters of 3 to 30 oocytes) in the tube-dwelling priapulid worm Paraselkirkia sinica from the Cambrian Stage 3 Xiaoshiba Lagerstätte (ca. 514 Ma, South China). These oocytes were accommodated within paired tubular ovaries located in the posterior half of the primary body cavity as in modern meiobenthic priapulid worms, thus indicating that the general organization of female tubular gonads in priapulid worms has remained virtually unchanged for half a billion years. Our findings provide for the first time, key information on the reproductive organs and strategies of early ecdysozoans, a huge animal clade that dominated Cambrian marine ecosystems and accounts for a large part of today's biodiversity (e.g. arthropods). Moreover, we also emphasize the critical role of ecology on the reproductive strategies and lifestyles of both modern and Cambrian worms.

Soft part preservation in hyolithids from the lower Cambrian (Stage 4) Guanshan Biota of South China and its implications

Hyoliths, known globally from the Cambrian to the Permian are generally recognized as early lophotrochozoan animals of uncertain phylogenetic position. The calcareous exoskeleton of hyolithids, one of the two subgroups of hyoliths, consist of four elements including a conical conch, an external operculum, and a pair of curved enigmatic helens, but their ecology and internal morphology have long been considered controversial. Here we present for the first time, remarkably preserved hyolithids with helens and interior soft tissues from the Guanshan Biota (Cambrian, Stage 4) of Shijiangjun section, Wuding County in eastern Yunnan Province of South China. Preserved soft tissues and organs encompass the digestive tract, muscle scars and connective tissues between the operculum and conch. Three modes of soft part preservation in hyolithids from the Guanshan Biota are described: (1) preservation through pyritization, (2) sediment-infilling of guts, and (3) bacterial biofilm pseudomorphs (expressed as large internal hollow cavities), resulting from endogenous bacterial decay of soft parts. Such differential preservation of hyolith guts indicates different taphonomic modes for the soft parts of Cambrian animals in the Guanshan Biota. The newly collected material yields the oldest preserved guts in hyolithids from Konservat-biotas, and reveals new evidence for reconstructing hyolithid anatomy, such as the articulation mechanism of the opercula and the internal structures in the conch. Comparisons with younger well-preserved hyolithid specimens indicate that the gut anatomy of hyolithids was evolutionarily conservative from the early Cambrian through to at least the Ordovician.

Symbiotic fouling of Vetulicola, an early Cambrian nektonic animal

Here, we report the earliest fossil record to our knowledge of surface fouling by aggregates of small vermiform, encrusting and annulated tubular organisms associated with a mobile, nektonic host, the enigmatic Cambrian animal Vetulicola. Our material is from the exceptionally preserved early Cambrian (Epoch 2, Age 3), Chengjiang biota of Yunnan Province, southwest China, a circa 518 million-year old marine deposit. Our data show that symbiotic fouling relationships between species formed a component of the diversification of animal-rich ecosystems near the beginning of the Phanerozoic Eon, suggesting an early escalation of intimate ecologies as part of the Cambrian animal radiation.