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.     

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.     

Evolution of feeding structure plasticity in marine invertebrate larvae: a possible trade-off between arm length and stomach size

The ability of an organism to alter its morphology in response to environmental conditions (phenotypic plasticity) occurs in several species of marine invertebrates. Examples are sea urchin and sand dollar larvae (plutei). When food is scarce, plutei produce longer food-gathering structures (larval arms and a ciliary band) and smaller stomachs than when food is abundant. However, it is unclear whether stomach size is actually induced through changes in morphogenesis or simply by food distending the stomach. Distinguishing between these two hypotheses is possible because plutei morphologically respond to food concentrations and change the length of their food-gathering structures before they are capable of feeding. More importantly, these two hypotheses provide insights to whether a trade-off exists between the response in food-gathering structures and the response in stomach size—a possible explanation for the evolution of feeding-structure plasticity in marine invertebrate larvae. In this study, I investigated whether sea urchin larvae (Strongylocentrotus purpuratus and S. franciscanus) reared in different amounts of food produced stomachs of different sizes before they were capable of feeding. Prior to having the ability to ingest food, larvae produced larger stomachs and shorter arms when food was abundant than when food was scarce, consistent with the hypothesis that food induced changes in morphogenesis. In addition, there was a strong negative correlation between the magnitude of plasticity in larval arm length and the magnitude of plasticity in stomach size. These results are consistent with the idea that a trade-off exists between the response in arm length and the response in stomach size, and at least in part, explains the evolution of feeding structure plasticity in plutei. This may also explain why feeding-structure plasticity has evolved in larvae of other taxa (e.g. other echinoderms and gastropods).     

Trusheimichnus New Ichnogenus From the Middle Triassic of the Germanic Basin,Southern Germany

The study of Jurassic Zoophycos from south‐eastern France has led us to some original results concerning its morphological organization and paleoenvironmental significance. Zoophycos represents a spreite constituted by a coiling lamina, constructed upwards in sediments, with only one opening at the sediment‐water interface. Zoophycos, produced by a deep sediment feeder, is emplaced late in an almost coherent substrate, intermediate between soft and firm ground. This suggests periods of very low or no sedimentation during which time the trace was constructed.

The Zoophycos studied developed on slope areas, characterized by a particular style of sedimentation: periods of nutrient supplies coming from proximal areas, alternating with periods of sedimentary breaks. Other paleoecologic and paleoethologic characteristics of the Zoophycos‐producer are discussed here: working activity, competition behavior, substrate modification, growing pattern and burrow oxygenation are the most interesting.     

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

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

Paleoecology of the marine endobenthos

Endobenthic animals, which reside within the sea bottom, include stationary suspension feeders, mobile deposit feeders and both stationary and mobile carnivores. Their activities, especially with regard to dwelling, feeding, walking/crawling and resting/nesting, are recorded as trace fossils.Abundance, diversity and density of some kinds of trace fossils allow interpretation of the population strategies of the trace-makers in terms of opportunistic (r-selected) and equilibrium (K-selected) strategies. Opportunistic ichnotaxa tend to be faciesbreaking traces, which are highly localized in low-diversity, high-density trace fossil associations in rocks representing environmental extremes (e.g., variable salinities, harsh temperatures, low oxygen levels or shifting substrates). Equilibrium ichnotaxa usually are restricted to particular sedimentary facies and are characteristic of high-diversity, low-dominance trace fossil associations in sediments reflecting stable, predictable environmental conditions.The most important environmental factors influencing the composition of trace fossil assemblages in marine settings are bathymetry, substrate, oxygen and hydrodynamic energy. The four factors are closely interrelated, because as water depth increases, there is a general decrease in sediment grain size and hydrodynamic energy of the depositional environment. As depth below the water—sediment interface increases, the firmness of the sediment (due to compaction and dewatering) increases and the oxygen content of interstitial waters drops drastically.Marine ichnofacies are largely substrate-controlled. Soupgrounds are water-saturated, incompetent substrates typified by highly compressed and usually unidentifiable burrows. Softgrounds commonly contain numerous distinctive burrows and are zoned bathymetrically by the Skolithos, Cruziana, Zoophycos and Nereites Ichnofacies. Firmgrounds are characterized by stiff, compacted sediments, in which traces of the Glossifungites Ichnofacies are excavated. Hardgrounds are cemented substrates, in which bioerosion traces of the Trypanites Ichnofacies are bored. Woodgrounds are woody materials that have been exposed to the sea and bored by bivalves, which produce characteristic traces of the Teredolites Ichnofacies. Tiering of endobenthic communities is common and is related to substrate preference of the burrowers and oxygen stratification of interstitial waters.     

Caging experiment in the deep sea: Efficiency and artefacts from a case study at the Arctic long-term observatory HAUSGARTEN

Present theories of deep-sea community organization recognize the importance of small-scale biological disturbances, originated partly from the activities of epibenthic megafaunal organisms, in maintaining high benthic biodiversity in the deep sea. However, due to technical difficulties, in situ experimental studies to test hypotheses in the deep sea are lacking. The objective of the present study was to evaluate the potential of cages as tools for studying the importance of epibenthic megafauna for deep-sea benthic communities. Using the deep-diving Remotely Operated Vehicle (ROV) “VICTOR 6000”, six experimental cages were deployed at the sea floor at 2500 m water depth and sampled after 2 years (2y) and 4 years (4y) for a variety of sediment parameters in order to test for caging artefacts. Photo and video footage from both experiments showed that the cages were efficient at excluding the targeted fauna. The cage also proved to be appropriate to deep-sea studies considering the fact that there was no fouling on the cages and no evidence of any organism establishing residence on or adjacent to it. Environmental changes inside the cages were dependent on the experimental period analysed. In the 4y experiment, chlorophyll a concentrations were higher in the uppermost centimeter of sediment inside cages whereas in the 2y experiment, it did not differ between inside and outside. Although the cages caused some changes to the sedimentary regime, they are relatively minor compared to similar studies in shallow water. The only parameter that was significantly higher under cages at both experiments was the concentration of phaeopigments. Since the epibenthic megafauna at our study site can potentially affect phytodetritus distribution and availability at the seafloor (e.g. via consumption, disaggregation and burial), we suggest that their exclusion was, at least in part, responsible for the increases in pigment concentrations. Cages might be suitable tools to study the long-term effects of disturbances caused by megafaunal organisms on the diversity and community structure of smaller-sized organisms in the deep sea, although further work employing partial cage controls, greater replication, and evaluating faunal components will be essential to unequivocally establish their utility.     

Role of reef fauna in sediment transport and distribution

Summary 1. Reef organisms may play a major role in the transport and distribution of sediment on the sea floor adjacent to coral reefs.2. Some fish such asMalacanthus plumieri (Bloch) selectively transport and collect certain types of sediment (such as larger coral and shell fragments).3. The random movement of crawling or burrowing organisms may cause a large amount of sediment to be shifted laterally on the sea floor. On slopes, a net downhill displacement may result.4. The surface configuration and internal structure of the sediment is rapidly changed by faunal mixing. Ripple marks formed by waves or currents are obliterated by the activity of organisms in only a few weeks in the environment studied. Internal structure (bedding) near the sediment-water interface is similarly destroyed in a short period of time.5. Larger clasts (including empty shells) on the sea floor tend to be buried by faunal undermining. The rate of burial depends primarily on the grain size of the substrate.6. The random movement of fauna on the sea floor may produce a predominantly concave-up orientation of pelecypod shells and shell fragments on the sea floor — the opposite of that produced by the activity of waves or currents.

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Die Rolle der Riffauna beim Transport und der Verteilung des Sediments. Untersuchungen von Tektite I und II

Kurzfassung Die Tierwelt scheint einen größeren Einfluß auf die Verfrachtung und Verteilung von Sedimenten in der Umgebung von Korallenriffen auszuüben als physikalische Vorgänge. Versuche, die während der Man-in-the-sea Projekte Tektite I und II bei St. John, U.S. Virgin Islands (Karibische See), durchgeführt wurden, galten unter anderem Studien über das Ausmaß des Transportes von Sedimentbestandteilen durch Fische. Ferner wurde die Geschwindigkeit, mit der Rippeln und Schichtungen durch tierische Wühlvorgänge zerstört und mit der Steine und andere Substratteile durch das Unterwühlen von Tieren umgelagert werden, untersucht. Ergänzende Beobachtungen wurden über die Orientierung leerer Muschelschalen angestellt, die je nachdem, ob sie von Bodentieren oder durch Wellen und Strömungen bewegt werden, entweder mit der konkaven Seite bevorzugt nach oben oder nach unten gelagert werden. Verschiedene Probleme der Beziehungen zwischen Riffauna und Sediment werden diskutiert.

     

The effects of sediment type on growth rate and shell allometry in the soft shelled clam Mya arenaria L

Hatchery-reared juvenile Mya arenaria L. were grown for 11 wk in replicated gravel, sand, mud, and pearl net treatments under flow-through sea-water conditions in Maine. Analyses of variance showed significant differences between sediment treatments for final shell length, dry meat weight, chondrophore growth increment, and percent shell weight. Growth of juvenile M. arenaria was more rapid in fine sediments than in coarse sediments or nets.Regression slopes of shell length-shell height and shell length-shell depth varied significantly between sediment treatments. Slower-growing clams from nets and gravel were more globose than clams from sand or mud treatments. Clams grown in sand were longer and narrower than those from mud. Differences in growth rates and shell form were attributed primarily to the physical properties of the substrata, and their effects on the scope for growth of M. arenaria.     

Diversity,Abundance, and Spatial Distribution of Ammonia-Oxidizing β-Proteobacteria in Sediments from Changjiang Estuary and Its Adjacent Area in East China Sea

Changjiang Estuary, the largest estuary in China, encompasses a wide range of nutrient loading and trophic levels from the rivers to the sea, providing an ideal natural environment to explore relationships between functional diversity, physical/chemical complexity, and ecosystem function. In this study, molecular biological techniques were used to analyze the community structure and diversity of ammonia-oxidizing bacteria (AOB) in the sediments of Changjiang Estuary and its adjacent waters in East China Sea. The amoA gene (encoding ammonia monooxygenase subunit A) libraries analysis revealed extensive diversity within the β-Proteobacteria group of AOB, which were grouped into Nitrosospira-like and Nitrosomonas-like lineages. The majority of amoA gene sequences fell within Nitrosospira-like clade, and only a few sequences were clustered with the Nitrosomonas-like clade, indicating that Nitrosospira-like lineage may be more adaptable than Nitrosomonas-like lineage in this area. Multivariate statistical analysis indicated that the spatial distribution of the sedimentary β-Proteobacterial amoA genotype assemblages correlated significantly with nitrate, nitrite, and salinity. The vertical profile of amoA gene copies in gravity cores showed that intense sediment resuspension led to a deeper mixing layer. The horizontal distribution pattern of amoA gene copies was nearly correlated with the clayey mud belt in Changjiang Estuary and its adjacent area in East China Sea, where higher β-Proteobacteria phylogenetic diversity was observed. Meanwhile, those areas with high amoA copies in the surface sediments nearly matched those with low concentrations of dissolved oxygen and ammonium in the bottom water.     

The diet of the Harlequin crab Lissocarcinus orbicularis,an obligate symbiont of sea cucumbers (holothuroids) belonging to the genera Thelenota,Bohadschia and Holothuria

The present paper characterizes, for the first time, the diet of the Harlequin crab Lissocarcinus orbicularis, an obligate symbiotic crab that associates with sea cucumbers (holothuroids) belonging to the genera Thelenota, Bohadschia and Holothuria. These tropical holothuroids host a rich symbiotic community in the Indo-West Pacific Ocean of which the Harlequin crab is the best known. The diet of L. orbicularis was characterized by analyzing the microscopic, molecular and isotopic signatures obtained from its gastric content. The presence of sea cucumber ossicles in the gastric mills of the crabs suggests that symbionts eat the superficial integument of their host and this was supported by the fact that Holothuroid DNA was detected in the stomach of L. orbicularis after DGGE and sequencing of the 18S rDNA gene. The stable isotopic δ¹³C and δ¹⁵N values of crab tissues were compared with diverse potential food sources including three holothuroids, three algae, one sea grass as well as the organic matter contained in the water column, in the sediment, and the second most abundant symbiont, the polychaete Gastrolepidia clavigera. The low δ¹⁵N values of crabs suggests that the crabs do not exclusively feed on sea cucumber tissue but assimilate diverse food sources such as sea grasses and organic matter contained in sediment that have similar δ¹³C values. There were no differences between the feeding of males and females but there was a positive correlation between the carapace length and the stable isotopic values indicating a shift of the food source as crabs grow larger.     

Changes in Ecosystem Function Across Sedimentary Gradients in Estuaries

The input of terrestrial silt and clay (hereafter mud) into coastal environments can alter sediment grain size distribution affecting the structure and functioning of benthic communities. The relationship between sediment mud content and macrofaunal community structure has been well documented, but not the effects on ecosystem function. In 143 plots from the mid-intertidal sites in 9 estuaries, we measured sediment properties, macrofaunal community composition and fluxes of O₂ and NH₄ ⁺ across the sediment–water interface to derive process-based measures of ecosystem function across the sand–mud gradient. We observed reductions in measures of macrofaunal diversity and decreases in the maximum density of key bioturbating bivalves (Austrovenus stutchburyi and Macomona liliana) with increased mud content. Concurrently, the maximum rates of sediment oxygen consumption (SOC), NH₄ ⁺ efflux (NH₄ ⁺) and biomass standardized gross primary production (GPP_Chl-a ) also decreased with increasing mud content. Environmental predictors explained 34–39% (P = 0.005–0.01) of the total variation in ecosystem function in distance-based linear models. After partitioning out the effect of mud, A. stutchburyi abundance was positively correlated and explained 25 and 23% (P = 0.0001) of the variation of SOC and NH₄ ⁺, respectively. Also, mud content (negatively correlated) and temperature (positively correlated) explained 26% of variability in GPP_Chl-a (P = 0.0001). Our results highlight the importance of increased mud content and the associated reduction in the abundance of strongly interacting key species on the loss of ecosystem function in intertidal sand flats.     

Lebensspuren von Meersauriern und ihren Beutetieren im mittleren Jura (Callovien) von Liesberg, Schweiz

Summary Numerous gutter-like furrows, up to 60 cm wide and up to 9 m long are preserved at the interface “Macrocephalus Beds”/“Callovian Marl” over a surface of 20 by 200 m. They are interpreted as feeding traces made by large marine vertebrates, most likely plesiosaurs and ichthyosaurs searching for food in the lime mud of the shallow Middle Jurassic sea floor. Possible prey animals were infaunal invertebrates (crustaceans) which produced an intricate meshwork of burrows (mainlyRhizocorallium irregulare andThalassinoides) in the bottom sediments, as well as infaunal bivalves. Evidence from cololites of predatory pelagic reptiles (ichthyosaurs, plesiosaurs) as well as reptile regurgitalites indicate that these animals fed not only on fast-swimming vertebrates and cephalopods but also on epi- and endobenthic invertebrates. In addition, the cololites show that the predators ingested considerable amounts of bottom sediment. Different sizes and shapes of the traces suggest that the gutters were produced by different reptiles or age groups. Candidates for the widest gutters are pliosaurs. Of the marine vertebrates known from Jurassic time, only the snout of adult pliosaurs of the genusLiopleurodon was broad enough to produce gutters more than 40 cm wide. Smaller, less than 15 cm wide gutters, could have been made by plesiosauroids or by the narrow pointed snouts of ichthyosaurs. Almost identical traces described from the Oxfordian of Spain and similar but smaller traces from the Lower Devonian of Prague are equally interpreted as feeding traces on the sea floor. Feeding traces of vertebrates in bottom sediments may give detailed information on the hunting behaviour of the predators. However, the attribution of the traces to definite vertebrate taxa remains uncertain.

     

Thiomicrospira pelophila,gen. n., sp. n., a new obligately chemolithotrophic colourless sulfur bacterium

From marine mud flats a very thin, comma- or spiral-shaped bacterium was isolated. The new organism was an obligately chemolithotrophic sulfur bacterium. Its physiology was found to be essentially similar to that ofThiobacillus thioparus. Because of the spirillum-like appearance it was proposed to classify this bacterium into a new genusThiomicrospira, with the species nameTms. pelophila. Tms. pelophila and a marineT. thioparus, which was isolated from the same mud, occupy different niches in this habitat.Tms. pelophila has a remarkable sulfide-tolerance as compared withT. thioparus. This property could be used for the specific enrichment ofTms. pelophila. The organism was also readily isolated in pure culture by filtering mud suspensions through a 0.22 Μm membrane filter.     

Channel country fluvial sands and associated facies of central-eastern Australia: Modern analogues of mesozoic desert sands of south America

The Channel Country, a region of wide fluvial plains criss-crossed by a reticulate pattern of anastomosing channels, and the adjacent sand dunes and clay pans of the Lake Eyre drainage basin occupy an area of 1.3 × 10⁶ km² of internal drainage in the arid east-central part of Australia. Beneath a surface of skin of mud, the sediment of the Channel Country is sand and some mud in the floodplain, as well as in levees and channels. The surface mud represents the overbank deposits of meandering channels that are superimposed on sheet sands of a relict braided system.With the other sediments (aeolian sand and lacustrine mud) of the Lake Eyre drainage basin, the Channel Country sediments represent the latest phase of deposition of the Cainozoic Birdsville Basin, which was initiated as an interior basin behind the dismembered rifted arch of the divergent Pacific margin. In their depositional and tectonic setting, the arid sediments of central-eastern Australia are modern analogues of the Mesozoic desert sands and other non-marine sediments that were deposited behind the pre-breakup arch and post-breakup half-arch of South America. The preceding Mesozoic Great Artesian Basin of central-eastern Australia contains volcanogenic sediment and was covered for a short time by an epeiric sea; its deposition was influenced by an uplift along the convergent Pacific margin, and it is analogous to other Gondwana basins in Antarctica, southern Africa, and South America that were yoked to the convergent Pacific margin.     

Using sediment alginate amendment as a tool in the restoration of Spartina alterniflora marsh

Growth and survival of transplanted vegetation for salt marsh restoration can be greatly affected by edaphic conditions. We investigated the potential for an organic carbon sediment amendment, alginate, to enhance establishment and growth of Spartina alterniflora transplants, as well as colonization by macroinvertebrates commonly associated with S. alterniflora (Uca sp. and Littoraria irrorata). Plots were established in a bare mud area previously inhabited by S. alterniflora, and transplant rhizospheres were either treated with alginate or left unamended as controls. Growth of transplanted S. alterniflora and colonization by Uca sp. and L. irrorata were evaluated over portions of three growing seasons. Within the first month post-transplantation, a nonsignificant trend toward greater live stem density emerged, but disappeared completely by week ten. Both Uca sp. and L. irrorata densities were significantly greater in the alginate treatment compared to the control only during the first ten weeks post-transplantation. However, the initial organic content of the sediment in the transplant area was more than twice that in the adjacent natural marsh. Therefore, our findings suggest that a threshold level of sediment organic matter above which alginate addition does not confer an advantage on transplanted S. alterniflora may exist, and the conditions under which amendments are most beneficial to transplanted S. alterniflora should be determined.     

Context-Specific Bioturbation Mediates Changes to Ecosystem Functioning

Species are often grouped according to their biological or functional traits to better understand their contribution to ecosystem functioning. However, it is becoming clear that a single species can perform different roles in different habitats. Austrohelice crassa, a burrow-building mud crab shifts its primary bioturbational role to that of a vertical mixer in non-cohesive sediments as frequent burrow collapse greatly enhances sediment reworking. We conducted in situ crab density manipulations in two sediment environments (a non-cohesive sand and a cohesive muddy-sand) to examine if the context-specific functional roles were linked to changes in solute fluxes across the sediment–water interface. Across both habitats, we show that A. crassa regulated nutrient cycling, creating strong density driven effects on solute exchanges. Increasing crab density increased sediment O₂ demand and the flux of NH₄ ⁺ from the sediment, indicating much of the response was physiologically driven. Clear interactions between A. crassa and microphytobenthos were also detected in both habitats. Despite lowering microphyte standing stock through deposit feeding, A. crassa increased benthic primary production per unit of chlorophyll a. Our experiment also revealed important context-specific differences, most notably for NH₄ ⁺ fluxes, which were higher where burrows and their associated microbial communities were most stable (muddy-sand). This study highlights the need to integrate interactions between organism behavior and habitat type into functional group studies to broaden conceptual frameworks and avoid oversimplification of highly complex organism–sediment interactions.     

Do biogenic sulphur compounds in phytodetritus act as cues for deposit-feeder activity? Field experiments and observations on the echiuran worm Maxmuelleria lankesteri

In temperate coastal seas, phytodetritus settling from the spring phytoplankton bloom is a potential food source for benthic deposit-feeders. The ability to exploit this seasonally variable resource could be enhanced by sensitivity to chemical cues signalling its arrival at the seabed. The biogenic sulphur compound dimethylsulphide (DMS), a breakdown product of dimethylsulphonioproprionate (DMSP) produced by some phytoplankton species, is a potential candidate for this role. We investigated the behavioural response of a sedentary surface deposit-feeder, the echiuran worm Maxmuelleria lankesteri, to DMS by observations and manipulative experimentation under natural conditions in a Scottish sea loch. Experimental addition of sediment enriched with DMSP-producing phytoplankton caused no significant increase in either the frequency of feeding by M. lankesteri or the rate of sediment ejection from observed burrows. Naturally occurring (DMSP+DMS) content of surface sediment was low during the winter, then peaked in April before declining in May. There was no consistent relationship between this parameter and rate of sediment ejection from M. lankesteri burrows. The results therefore provide no evidence that M. lankesteri uses DMSP or DMS as a stimulus to increased activity. An observed imbalance between the frequency of surface deposit-feeding and sediment ejection from individual burrows remains unexplained.     

An additional example of the ichnogenus Zoophycos in Cambrian strata (Gallatin Formation; Wyoming,USA)

The ichnogenus Zoophycos is relatively rare in Cambrian-age sedimentary rocks, being far more commonly encountered in post-Devonian, and particularly post-Paleozoic, strata. A new occurrence of Zoophycos from the Upper Cambrian Gallatin Formation of the Bighorn Mountains, north-central Wyoming, USA, is reported herein. Material consists of a single specimen preserved within quartz silt representing the upper surface of a graded quartz arenite bed. The burrow is a relatively wide, curved, horizontally oriented lobe with a poorly visible marginal tube but well-developed internal lamellae. Discovery of this specimen contributes to the paleobiogeographic distribution of lower Paleozoic Zoophycos, adds to the literature suggesting that most Cambrian Zoophycos were horizontal and shallow-tier structures, and provides further support to the notion that relatively complex deposit-feeding behavior evolved earlier in the Phanerozoic than had long been recognized.     

A comparative study on the tubes and feeding behaviour of eight species of corophioid Amphipoda and their bearing on phylogenetic relationships within the Corophioidea

Observations are presented on mouthpart functional morphology, and on feeding, grooming and defaecatory behaviour of eight species of corophioid Amphipoda, viz. Corophium bonnellii, Lembos websteri, Aora gracilis, A. spinicornis, Gammaropsis nitida, Ericthonius punctatus, Jassa falcata and J. marmorata. These data are considered in relation to tube structure and amphipod posture in relation to the tube. All these species occupy double-ended cylindreical tubes made from ''amphipod silk'' secreted by the third and fourth preraeopods, incorporating sediment and other debris to varying degrees. Uniquely among this set of species, however, E. punctatus has a tube that is architecturally distinct. It tapers along its length and has a distinctive oblique main entrance at its widest end. This end is used preferentially. The other species studied use either opening with equal facility. Such a feature is adaptive in facilitating deployment of the antennae and shielding the head of E. punctatus.Two groupings of species are propose: group A which feed inside their tube using pleopod-induced through-tube currents, and group B which feed outside or at the entrance to their tube using external water currents. Group A includes C. bonnellii, L. websteri and the Aora species. Group B includes E. punctatus and the Jassa species. Gammaropsis nitida exhibits traits from both groups, adding weight to its perceived status as a genus representative of the stem corophioid. The ischyrocerid habit of externalizing food-gathering may be regarded as the first step along an evolutionary line leading to the rod-building podocerid types and ultimately towards the caprellids. All species examined show a degree of flexibility in their feeding habits which helps to explain the success of this taxon, which has radiated into a great diversity of aquatic biotopes.