Ethological analysis of the trace fossil Zoophycos: hints from the Arctic Ocean

Löwemark, L. 2011: Ethological analysis of the trace fossil Zoophycos: Hints from the Arctic Ocean. Lethaia, Vol. 45, pp. 290–298. The distribution of the trace fossil Zoophycos in Quaternary marine sediments from the Arctic Ocean was studied in twelve piston and gravity cores retrieved during the Swedish icebreaker expeditions YMER80, Arctic Ocean‐96 and LOMROG I & II. The sampled cores span an area from the Makarov Basin to the Fram Strait. Zoophycos was only found in two cores taken at more than 2 km water depth on the slope of the Lomonosov Ridge, but was absent in cores obtained at shallower depth, confirming earlier observations of the trace maker’s bathymetric preferences. The two cores containing Zoophycos are characterized by quiet sedimentation and slightly enhanced food flux compared with the general Arctic. The occurrence of Zoophycos in these cores in a setting that is characterized by extreme seasonal variations in food flux due to the total ice coverage during winters and high primary productivity during the long summer days, is interpreted to be a cache‐behaviour response to pulsed flux of food to the benthic realm. □Arctic Ocean, ethology, Quaternary, spreiten, trace fossils, Zoophycos.     

Zoophycos composite ichnofabrics and tiers from the Permian neritic facies in South China and south‐eastern Australia

Gong, Y.‐M., Shi, G.R., Zhang, L.‐J. & Weldon, E.A. 2009: Zoophycos composite ichnofabrics and tiers from the Permian neritic facies in South China and south‐eastern Australia. Lethaia, Vol. 43, pp. 182–196. Zoophycos composite ichnofabrics (ZCI) comprising two or more suites of the same form of Zoophycos are widespread and densely distributed in Early and Middle Permian (Cisuralian–Guadalupian) neritic limestones (Qixia and Maokou Formations) of palaeotropical origin in the Laibin area, Guangxi, South China. Similar ZCI also occur in neritic greywackes of glaciomarine origin from the Middle Permian (Guadalupian) Westley Park Sandstone Member (Broughton Formation) in the southern Sydney Basin, south‐eastern Australia. Zoophycos from both regions consists of planar spreite with major and minor lamellae and a cylindrical tunnel interpreted as a marginal tube and/or axial shaft. The cylindrical tunnel is herein considered to be an essential component of Zoophycos, and thus can be used to define and characterize the morphological variability of Zoophycos. It is suggested that the variation of spreite and major and minor lamellae originated from the different morphologies and migration manners of the cylindrical tunnel. The shallowest, shallow, middle and deepest Zoophycos tiers have been distinguished in ZCI on the basis of cross‐cutting relationships, the soft‐sediment deformation and the contrast in colour between Zoophycos and its host rock. The multiple tiers may represent the substrate consistency spectrum from a softground through a stiffground to a firmground. The different Zoophycos tiers may have been constructed by tracemakers of either different or the same taxonomic affinities in response to the gradual accretion and lithification of sediment layers on the seafloor. The tracemakers appeared to be very sensitive to neither climate nor lithology. The width of the planar spreiten of Zoophycos decreases slightly with the depth of tiering in ZCI. □Composite ichnofabric, Permian, South China, south‐eastern Australia, tier, Zoophycos.     

Complex behavioural patterns and ethological analysis of the trace fossil Zoophycos: evidence from the Lower Devonian of South China

The trace fossil Zoophycos is abundant in the shallow‐marine deposits (tempestites) of the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by deposit‐feeding behaviour, where the animal constructed the trace upwards without leaving faeces in the spreiten. The width of the marginal tube in different whorls is almost constant. The scratches are observed on the wall of the marginal tubes. The Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos‐maker might have been a vermiform polychaete instead of a predator such as a decapod crustacean (Spongeliomorpha producer). Based on stratigraphical and ichnological features, the complex trace fossils resulted from the complex activity of different opportunistic organisms (r‐strategist) that quickly occupied and thrived within the quiet, nutrient‐rich environment after storm events.     

Evolutionary trend of Zoophycos morphotypes from the Upper Cretaceous–Lower Miocene in the type pelagic sections of Gubbio,Italy

About 200 Zoophycos specimens, including 90 specimens studied in detail, have been analysed in the continuous Upper Cretaceous–Lower Miocene pelagic sedimentary type sections of the Gubbio area (the Contessa Highway, Contessa Quarry and Bottaccione sections, Northern Apennines). The sediments are reddish to grey limestones and marls of the Scaglia Group and marls with volcaniclastic deposits of the Bisciaro Formation. The aim was to examine the evolutionary trend of what is probably the most debated trace fossil of all time, from the Upper Cretaceous to Lower Miocene. Despite having been found in beds ranging from the Cambrian to the present, no consensus has been reached regarding mode of construction, tracemaker or ethological explanation for Zoophycos. Four Zoophycos morphotypes are recognized at Gubbio showing variations of major and minor lamellae, apex, lobes and whorls: the Cretaceous–Eocene cone‐shaped type 1, the Upper Eocene–Middle Oligocene helicoidal type 2, the Oligocene lobate type 3 and the Upper Oligocene–Lower Miocene flat type 4. The very high ichnodensity in some beds (hundreds of specimens in discrete levels of the Bisciaro Formation, now destroyed by quarrying) seems to find explanation in abnormal concentrations of phytodetritus and organic matter on the seafloor in some periods. This very high abundance in discrete levels reflects a change in sedimentation and seafloor conditions at pre‐flysch deposition. Due to such high ichnodensity, many adjacent specimens display deformed outer margins. Taphonomic analysis shows a variation of whorls, laminae and U‐shaped lobes, reflecting ontogenetic development of the tracemaker(s) (?sipunculid worms).     

Zoophycos in storm-affected environments: a case study from lower Maastrichtian deposits of the Mateur-Beja area (Northern Tunisia)

Abstract

The lower Maastrichtian deposits of the Mateur-Beja area in northern Tunisia are mainly composed of fine-grained marl and limestone alternations occasionally interbedded by coarse-grained calcarenites and gravel deposits. In the coarse-grained intervals sedimentary structures are indicative of storm-induced high-energy currents in an outer ramp to slope setting and of local reworking by bottom currents in the basin. In deeper environments, fine-grained sediments accumulated mainly while settling from storm-induced suspensions. The lower Maastrichtian deposits contain abundant Zoophycos exhibiting two main morphotypes, skirt-shaped Zoophycos in deposits around storm-wave base and tongue-shaped Zoophycos in somewhat deeper sediments. The types differ in burrow architecture, morphology of lobes, and size of structural elements. These differences are attributed to different behavioral programs modulated by the availability of benthic food that decreased seaward. Storm-affected environments seem to be a prerequisite for these Zoophycos-producers to choose their habitat.     

First Record of Graphoglyptids in Cyprus: Indicative Presence of Turbidite Deposits at the Pakhna Formation

Abstract

Ichnological analysis at the Pakhna Formation (Miocene, Cyprus) reveals, for the first time, the presence of graphoglyptid structures. The Pakhna Formation is dominated by pelagic/hemipelagic sediments, together with contourite, reworked turbidite and turbidite facies. Thus, a complex interaction between pelagic, bottom-current and gravitational sedimentation is envisaged. The discernment of facies is sometimes difficult, based principally on stratigraphic and microfacies features. The record of Helminthorhaphe as a component of the graphoglytid group supports the presence of turbidites, facilitating their differentiation from the other facies. Nereites ichnofacies, punctuating a dominant/background Zoophycos ichnofacies, would reflect changes in paleoenvironmental conditions. This record offers significant perspectives to advance in the study of bottom-current and gravitational processes through outcrop examples.     

Rhizocorallium hamatum (Fischer-Ooster 1858), a Zoophycos-like trace fossil from deep-sea Cretaceous-Neogene sediments

Rhizocorallium hamatum (Fischer-Ooster 1858) is a trace fossil of the Zoophycos group, which is distinguished by its mostly horizontal, branched spreite lobes. It has so far, been ascribed mainly to Zoophycos, but the latter should be limited to forms having helical whorls, which are absent in R. hamatum. It has also been ascribed to Phycosiphon, which, however, shows J-shaped spreite lobes, while the lobes of R. hamatum are U-shaped. R. hamatum is very similar to R. commune var. irregulare, but the latter displays a distinctly wider marginal tunnel with respect to lobe width. R. hamatum occurs from the Turonian to Eocene, possibly from the Hauterivian to Oligocene, but mostly from Maastrichtian to Palaeocene, deep sea, mainly turbiditic sediments rich in marl. The tracemaker, probably a ‘worm’-like invertebrate, ingested an organic-rich mud of the background sediment and relocated it into the middle to deep tiers within the underlying turbiditic marl, mostly in form of faecal pellets packed within the spreite lobes, for further use as a food resource. This way of feeding was a response to food deficiency on the deep-sea floor.     

Responses to temperature,heavy metal and sediment stress by the giant clam Tridacna squamosa

Increasing seawater temperature, heavy metal concentration and sediment load, all represent environmental stress factors found in the Gulf of Thailand. In the present study we investigate and compare the physiological responses of exposure to increased water temperature (+ 3°C) and copper (50 μg Cu/l) by the giant clam Tridacna squamosa, and the behavioural response to sedimentation.

Both temperature and copper caused significant decrease in P_g: R ratios, although by different physiological responses. In clams exposed to copper, the decrease in P_g: R is caused by reduced photosynthesis, whereas the clams exposed to increased temperature maintained a high photosynthetic rate, but significantly increased their respiration.

Clams responded to additional sediment with increased activity. This augmented activity was further increased for clams previously stressed by copper.     

Anaerobic mineralization of marine sediment organic matter: Rates and the role of anaerobic processes in the oceanic carbon economy

Abstract

This paper addresses three related questions: (1) What factors control the efficiency of carbon burial in sediments? (2) Are rates of anaerobic organic matter degradation intrinsically lower than aerobic rates? (3) How important are anaerobic processes in the global marine sediment carbon economy?

Carbon burial efficiency (the ratio of the carbon burial rate and the carbon flux to the sediment surface) was estimated from literature data for a range of environments and was shown to be a function of sedimentation rate. No difference independent of sedimentation rate was found between aerobic and anaerobic sediments.

A review of recent microcosm and laboratory studies shows that anaerobic rates are not intrinsically lower than aerobic rates; fresh organic matter degrades at similar rates under oxic and anoxic conditions. Aerobic decomposition rates near the sediment surface are typically greater than anaerobic rates at depth because the most labile carbon is consumed before it can be buried in the anoxic zone.

A model approach was taken in estimating the importance of anaerobic processes in the global marine sediment economy, instead of extrapolating measured rates as done previously. The result, 150 Tg C yr^?1, is two to nine times lower than previous estimates. This rate is about 9% of the global aerobic carbon oxidation rate and is about equal to the rate of long‐term carbon burial. The importance of anaerobic processes in marine sediments lies in their role in determining the amount of carbon preserved, not in the amount of carbon remineralized overall.     

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

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

Alteration of burrow‐related behaviour of the norway lobster,nephrops norvegicus during infection by the parasitic dinoflagellate hematodinium

This study has used time‐lapse video recordings to analyze the burrow‐related behaviour of N. norvegicus under a 12 h light: dark regime in aquarium conditions, and compared this with the behaviour of lobsters infected by the parasitic dinoflagellate Hematodinium. Uninfected lobsters performed a higher number of burrow departures, of longer duration, during the hours of darkness than during the light. Infected lobsters performed a relatively constant number of departures of similar‐duration in the light and the dark periods. However, the mean number of departures per day performed by infected lobsters (70 day^‐1) was more than double that of uninfected lobsters (30.1 day^‐1), while the mean duration of burrow departures performed by infected lobsters (258.5s departure^‐1) was more than six times greater than in uninfected lobsters (38.7s ? departure^‐1). This led to a more than ten‐times increase (from 1.7 to 19.4%) in the percentage of the day spent out of the burrow by infected lobsters. This altered burrow‐related behaviour could be due to the nutritional demands of the parasites on infected lobsters, causing an increased requirement to forage, or alternatively to a ‘functional hypoxia’, due to the additional respiratory demands of the parasite, causing the host to emerge from the burrow onto the sediment surface to seek oxygen. Implications for the increased time spent out of the burrow are discussed in relation to availability of infected lobsters by trawlers and predators, and to the use of infection prevalence data from trawl‐caught samples in stock assessment models for N. norvegicus.     

Image processing techniques to improve characterization of composite ichnofabrics

Abstract

Image processing techniques, including the Analyze Particles tool offered by Fiji software and the Intensity Profile by ICY (IP-ICY), were applied in core and outcrop examples to improve characterization of autocomposite ichnofabrics. Analyze Particles gives information about particle shape and size in the studied image. This tool was applied to Chondrites assemblages in composite ichnofabrics in view of selected images of modern marine hemipelagic cores from Site U1385 of IODP Expedition 339. Differences in size, relative abundance, density and distribution of Chondrites were interpreted as related to variations within the population of Chondrites tracemakers. Intensity Profile quantifies pixel values of the infilling material of traces, proving helpful to discriminate between specimens, evaluate the horizon of colonization, and approach the penetration depth. Its application to the analysis of Zoophycos improves characterization of composite ichnofabrics from deep-sea pelagic calcilutites of the Petra Tou Romiou section (Eocene Lefkara Formation; southern Cyprus). Some suites of Zoophycos were interpreted as associated with different phases of colonization from several horizons. Moreover, it was possible to discriminate structures pertaining to different specimens, as opposed to those from the same specimen.     

Using repeated winter surveys to estimate changes in abundance of seed‐eating passerines

Capsule Winter Atlas surveys of 16 species on lowland farmland revealed significant changes in count for four species.

Aims To estimate changes in abundance between the early 1980s and late 1990s, of wintering seed‐eating passerines, in ‘core’ areas of lowland Scotland.

Methods Ninety‐five Scottish 10‐km squares were selected that held high numbers of seed‐eating passerines in the 1981–84 Atlas of Wintering Birds in Britain and Ireland. The same survey methods were used to resurvey these in winters 1997/98 and 1998/99, and visits were matched as closely as possible for duration and date. Analyses compared counts between the two survey periods for 16 species of seed‐eating passerines and, for 12 of these, differences were also compared with national breeding population trend information for the same period.

Results Mean Corn Bunting Emberiza calandra count per visit declined by 62% between the early 1980s and late 1990s, a difference which was statistically significant (P = 0.026). Significant increases were recorded for Yellowhammer Emberiza citrinella (up 62%), Common Linnet Carduelis cannabina (up 3.4‐fold) and European Goldfinch Carduelis carduelis (up 13‐fold). For 12 species for which national breeding population trend data were available, trends were weakly positively correlated (r _s = 0.43, P = 0.08) with those from our results, but several species trends were more positive in our study. This difference was particularly marked for Eurasian Tree Sparrow Passer montanus, Goldfinch and Linnet.

Conclusion Repeating Winter Atlas surveys offers a useful additional method for assessing population trends. They are particularly useful in a region with low observer coverage and for species that are poorly covered by long‐term bird monitoring data sets. It would be valuable to validate this approach at a regional level, especially in a region for which detailed long‐term bird monitoring data are available.     

Trace fossils and their environmental significance in Dinantian carbonates of Belgium

Dinantian carbonates of the Tournai and Dinant regions, southern Belgium, contain several trace fossil assemblages. These assemblages consist of the following ichnogenera:Cruziana, Rhizocorallium,Scalarituba, Scolicia, Skolithos, Phycosiphon, Planolites andZoophycos. Scalarituba andPhycosiphon are reported for the first time from the Carboniferous of Western Europe. The ichnofossils occur in discrete ichnofacies.Skolithos andZoophycos ichnofacies occur within the Ivorian (Upper Tournaisian). TheCruziana andSkolithos ichnofacies are found in the Moliniacian (Lower Visean). In the Dinant region, ichnofossils record a change from infauna to epifauna at the Ivorian-Moliniacian transition. Woodgrained chert is recognized and corresponds stratigraphically and palaeogeographically with those from the Western Interior United States.     

Ichnology of an ephemeral lacustrine/alluvial plain system: Jurassic East Berlin Formation,Hartford Basin,USA

A trace fossil assemblage from the Lower Jurassic East Berlin Formation of the Newark Supergroup, Hartford Basin, New England, USA, includes: Scoyenia gracilis, Skolithos ichnosp., Palaeophycus striatus, Planolites montanus, Fuersichnus ichnosp., fusiform burrows, pelleted material, an escape structure, and large burrows.

This assemblage is assigned to the Scoyenia ichnofacies. Specific lebensspuren are not limited to specific lithofacies; instead, their initial distribution seems to have been influenced principally by water availability within an ephemeral lacustrine/alluvial plain system. Other factors in distribution may have included amounts of organic matter, patterns of sedimentation, sediment grain size, biotic factors (settling from invertebrate drift, competition), and additional abiotic factors (wind deflation, waves, currents, desiccation, soft‐sediment deformation, evaporite formation, pedoturbation).

Extreme environmental conditions within the original depositional setting strongly influenced the availability of water which, in turn, strongly influenced the paleoecology of burrowing invertebrates in this nonmarine system.     

Shift of Bacterial Community Structures in Sediments From the Changjiang (Yangtze River) Estuary to the East China Sea Linked to Environmental Gradients

Abstract

The Changjiang estuary and its adjacent East China Sea (ECS) have been considered as one of the most dynamic areas significantly contributing to elemental exchanges globally. The purpose of this study was to understand the alteration of microbial consortia at the interface of riverine and coastal environments in relation to environmental variations as well as their roles in biogeochemical cycling at this dynamic region. We sampled surface sediment samples at 4 stations from the estuary to coastal regions of the ECS. Along with collections of physicochemical parameters, we sequenced bacterial 16S rRNA genes of clones from each sample. Results showed a distinct transition of bacterial community from typical freshwater sediment phyla (e.g., Betaproteobacteria and Firmicutes) to those commonly inhabited in saline environments (e.g., Deltaproteobacteria and Gammaproteobacteria). The bacterial group at the transition zone characterized by high accumulation of organic matters and intense mixing of riverine and coastal waters was most diverse. Bacterial community structures at two ECS stations showed a similar pattern but contained different dominant taxa, shifting from Deltaproteobacteria-affiliated sulfate-reducing bacteria at the station closer to the shore to Gammaproteobacteria-affiliated nitrate-reducing bacteria further offshore. It suggested that the sedimentary bacterial community structure was related to salinity, sediment type, and substrate availability and composition.     

Palaeoecology of solitary corals in soft-substrate habitats: the example of Cunnolites (upper Santonian, Eastern Alps)

The upper Santonian Hofergraben Member (Eastern Alps) provides an example of a soft‐substrate habitat suited mainly for solitary corals (Cunnolites), for colonial forms of solitary coral‐like shape (Placosmilia, Diploctenium), and for colonial corals of high sediment resistance (e.g. Actinacis, Pachygyra). The Hofergraben Member consists mainly of silty‐sandy marls of wave‐dominated, low‐energy shore zone to shallow neritic environments. Substrates of soft to firm mud supported level‐bottoms of non‐rudist bivalves, gastropods, solitary corals, colonial corals, rudists, echinoids, and benthic foraminifera. Boring and/or encrustation of fossils overall are scarce. In the marls, Cunnolites is common to abundant. Both a cupolate shape and a lightweight construction of the skeleton aided the coral to keep afloat soft substrata. Cunnolites taphocoenoses are strongly dominated by small specimens (about 1–3 cm in diameter). Cunnolites was immobile and mostly died early in life upon, either, smothering during high‐energy events, rapid sedimentation associated with river plumes, or by toppling and burial induced by burrowing. Comparatively few large survivor specimens may show overgrowth margins interpreted as records of partial mortality from episodic sedimentation or tilting on unstable substrate. Scattered pits and scalloped surfaces on large Cunnolites may have been produced, in some cases at least, by predators (durophagous fish?). Post‐mortem, large Cunnolites provided benthic islands to corals, epifaunal bivalves and bryozoans. In a single documented case of probable in vivo contact of Cunnolites with the colonial coral Actinastraea, the latter prevailed.     

Threshold determination for olfactory receptors of the spiny lobster

The shallow water caridean shrimps Heptacarpus pictus and H. paludicola are polymorphic in colour pattern. Populations of these species collected over colour‐variable substrates showed the greatest degree of coloration in terms of the proportion of individuals displaying a distinct colour pattern. The frequencies of H. Pictus colour morphs varied significantly between most sampling periods. Apostatic selection by fish predators is suggested as one hypothesis which could explain these changes in morph frequency.

Experiments with Heptacarpus pictus on rapid colour change showed that, although some pigment migration did occur in the chromatosomes studied, the macroscopic appearance of the colour patterns was not altered when shrimps were shifted from black to white backgrounds or vice versa. The results of background choice experiments with H. pictus suggest that these shrimp do not seek out colour backgrounds that would seem to be a matching background in a concealing coloration. This behaviour is consistent with the morphology of the colour patterns which appears to be a disruptive coloration rather than a concealing coloration that closely matches a particular substrate.     

Morphometry and sedimentation as regulating factors for nutrient recycling and trophic state in coastal waters

The aim of this work is to quantify the importance of morphometry and sedimentation/resuspension on nutrient recycling and trophic characteristics in coastal waters. Extensive field work has been carried out in 23 coastal areas in the Swedish and Finnish part of the Baltic Proper. Sediment traps were deployed for two one-week periods in all areas. On average, 56% of the total sedimentation in sediment traps 3 m below the water surface (SedS) and 62% of the total sedimentation on sediment traps 1 m above the bottom (SedB) was resuspended material. Coastal morphometric parameters, surface water retention time and bottom dynamic conditions were determined for all areas. There is a marked relationship between SedS and inorganic-N concentration in the surface water. The relationship was improved significantly by using sedimentation of the resuspended fraction at 3 m water depth (SedR) instead of SedS.This led to the hypothesis that increased concentration of inorganic nitrogen in the surface water results from increased mineralisation of resuspended organic particles. A model describing SedS is presented where inorganic nitrogen concentration, the water surface area and the surface water retention time can explain 82% of the variation in SedS. In another model inorganic nitrogen and water surface area can explain as much as 93% of the variation in SedR.These results emphasise the importance of resuspension for nutrient recycling and trophic state in coastal waters. The importance of coastal morphometry and surface water retention time on total sedimentation and nutrient recycling makes it possible to classify coastal areas in terms of potential nutrient recycling capacity/trophic state from these simple sensitivity parameters.     

Zoophycos and umbellula (Pennatulacea): Their synthesis and identity

The trace fossil Zoophycos has the shape of a low-angled helicoidal sheet constructed of numerous spirally radiating and adjacent dwelling burrows, each lined with retextured sediment derived from the adjacent or underlying beds. Such spiral sheets are here considered the products of a sea pen or closely allied organism with its trunk at the sea floor and its tubular stem suspended in, and voluting laterally through, the underlying soft sediments. Peristaltic movements of the stem fluidise the sediment in the annular space around the stem and allow both the lateral voluting motion of the stem and the pumping of nutrient-rich mud up to the sea floor. The organic content of the mud is delivered to the feeding filters of polyps of the animal which lie along the length of the stem and through these it passes into circulatory canals. Nutrient particles are then distributed by water currents through the length of the organism.The above proposed mechanism of feeding fits the probable ecology and structure of the Umbellulidae and explains the form, structure and occurrence of Zoophycos and related fossils.