Disturbance,emigration, and refugia: How the mud snail,Ilyanassa obsoleta (Say), affects the habitat distribution of an epifaunal amphipod,Microdeutopus gryllotalpa (Costa)

In the presence of the mud snail Ilyanassa obsoleta (Say), the tubicolous amphipod Microdeutopus gryllotalpa (Costa) emigrates to snail-free sediments, as demonstrated in laboratory and field experiments. Emigration occurs predominantly in the dark when the amphipod is most active. Unlike crevices, the thickness of sediments in which the amphipod is established offers no protection from snail disturbance. Emigration is shown to be caused by the disturbance generated by the snail's plowing and burrowing across the sediment surface, and not a response to a reduction in the shared microfloral food supply. The crawling and burrowing of the smaller mud snail, Hydrobia totteni Morrison, does not disturb Microdeutopus, supporting the hypothesis that relative body sizes affects the ability of bioturbators/burrowers to disturb tube-dwellers. As the burrows of Microdeutopus extend only ≈2 cm below the sediment surface, thick mud layers do not offer any refuge from Ilyanassa. However, very small solid surfaces (≈1–2 mm in relief) to which the amphipods build tubes do provide some protection from Ilyanassa. In soft-sediment benthic communities, such small structures may provide significant refuge for small epifauna and shallow burrowing infauna escaping from small-scale, biogenic disturbance.     

Habitat preferences of two estuarine burrowing crabs Helice crassa Dana (Grapsidae) and Macrophthalmus hirtipes (Jacquinot) (Ocypodidae)

The horizontal and vertical distributions of two species of endemic, burrowing mud crabs Helice crassa Dana 1851 (Grapsidae) and Macrophthalmus hirtipes (Jacquinot 1853) (Ocypodidae) are described for the Avon-Heathcote Estuary (43°33′S: 172°44′E), Christchurch, New Zealand. Substratum preference is shown to be the most important factor influencing mud crab distribution, but lack of tolerance to salinities below 4‰ is also a significant factor preventing M. hirtipes from occurring at points close to freshwater input. Both species had similar sediment organic content and particle size requirements. Helice crassa was concentrated in well-drained, compacted sediments above mid-tide level, whilst Macrophthalmus hirtipeswas found in waterlogged areas below mid-tide level. This vertical separation is shown not to be caused by differential desiccation tolerances, but by feeding and burrowing adaptations related to these different substrata.     

Cutting out the middle clam: lucinid endosymbiotic bacteria are also associated with seagrass roots worldwide

Seagrasses and lucinid bivalves inhabit highly reduced sediments with elevated sulphide concentrations. Lucinids house symbiotic bacteria (Ca. Thiodiazotropha) capable of oxidising sediment sulphide, and their presence in sediments has been proposed to promote seagrass growth by decreasing otherwise phytotoxic sulphide levels. However, vast and productive seagrass meadows are present in ecosystems where lucinids do not occur. Hence, we hypothesised that seagrasses themselves host these sulphur-oxidising Ca. Thiodiazotropha that could aid their survival when lucinids are absent. We analysed newly generated and publicly available 16S rRNA gene sequences from seagrass roots and sediments across 14 seagrass species and 10 countries and found that persistent and colonising seagrasses across the world harbour sulphur-oxidising Ca. Thiodiazotropha, regardless of the presence of lucinids. We used fluorescence in situ hybridisation to visually confirm the presence of Ca. Thiodiazotropha on roots of Halophila ovalis, a colonising seagrass species with wide geographical, water depth range, and sedimentary sulphide concentrations. We provide the first evidence that Ca. Thiodiazotropha are commonly present on seagrass roots, providing another mechanism for seagrasses to alleviate sulphide stress globally.Subject terms: Microbial ecology, Plant ecology, Soil microbiology     

Evidences of habitat displacement between two common soft-bottom SW Atlantic intertidal crabs

The intertidal burrowing crab Chasmagnathus granulatus Dana is the dominant species in soft sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 41°S) where it produces dense and extensive burrowing beds. The mud crab Cyrtograpsus angulatus Dana coexists with Ch. granulatus in this area, but it also inhabits areas to the south (northern and central Argentinean Patagonia). A survey covering both areas showed that C. angulatus rarely live in burrows when coexisting with Ch. granulatus, but form large burrowing beds when not coexisting with Ch. granulatus. When both species coexisted, burrowing beds of C. angulatus are restricted to sandy-muddy areas. Only rarely are burrows of C. angulatus found within Ch. granulatus beds. However, when Ch. granulatus were experimentally excluded within their burrowing beds, new settlers of C. angulatus made burrows and maintained them until they reached large size. Paired (inside and outside Ch. granulatus burrowing bed) sampling during high tide using beach nets showed that C. angulatus rarely venture inside the Ch. granulatus crab beds. Other field experiments showed that adults Ch. granulatus always displace C. angulatus from burrows. Furthermore, in several sites located south of the limit of distribution of Ch. granulatus at the Patagonian coast, soft bare intertidals are dominated by burrowing beds of C. angulatus mixed with the congener C. altimanus Dana. Together, these evidences suggest that the mud crab C. angulatus is displaced from soft bottom areas by the burrowing crab Ch. granulatus. It is an example of competitive exclusion through aggressive interference in soft-bottom habitats when the shared resource is the access to sediment surface, a two-dimensional well-defined resource.     

The effect of natural populations of the burrowing and grazing soldier crab (Mictyris longicarpus) on sediment irrigation, benthic metabolism and nitrogen fluxes

The aim of this study was to determine the effect of sediment grazing and burrowing activities of natural populations of Mictyris longicarpus on benthic metabolism, nitrogen flux and irrigation rates by comparing sediments taken from minimum disturbance exclusion cages and adjacent sediments subject to M. longicarpus activities. M. longicarpus reduced sediment surface chlorophyll a (approximately 77%), organic carbon (approximately 95%) and total nitrogen concentrations (approximately 99%) in comparison to ungrazed sediments. Consequently, they significantly reduced gross benthic O₂ production (about 71%) and sediment O₂ consumption (approximately 46%). Mean N₂ fluxes showed net effluxes (276-430 μmol m⁻² day⁻¹) in the presences of M. longicarpus and net uptakes (194.09-449.21 μmol m⁻² day⁻¹) where they were excluded. The net uptake of N₂ was most likely due to cyanobacteria fixing of N₂, as dense microbial mats became established over the sediment surface in the absence of M. longicarpus grazing activity. Sediment irrigation/transport rates calculated from CsCl tracer dilution indicated greater irrigation rates in the exclusions (12.12-16.22 l m⁻² h⁻¹) compared to inhabited sediments (6.33-11.73 l m⁻² h⁻¹) and this was again was most likely due to the lack of grazing pressure which allowed large populations of small burrowing polychaetes to inhabit the organic matter rich exclusion sediments. As such, the main influence of M. longicarpus was the interception and consumption of transported organic material, benthic microalgae and other small infaunal organisms resulting in the removal of approximately 0.06 g m⁻² day⁻¹ of nitrogen and 12.12 g m⁻² day⁻¹ of organic carbon. This “cleansing” of the sediments reduced sediment metabolism and the flux of solutes across the sediment water interface and ultimately the heavy predation of M. longicarpus by transient species such as stingrays, results in a net loss of carbon and nitrogen from the system.     

The utilization of fly ash by two tube-building polychaetes

Benthic tube-building polychaetes which utilize sediment particles have been found to incorporate fly ash in their tubes. Experiments were conducted on North Sea specimens of Myriochele oculata Zachs and Pectinaria koreni (Malmgren) in artificial sediments offering a range of fly ash concentrations. A marked selection for non-ash particles was shown by Myriochele oculata, and a marginal selection against ash by Pectinaria koreni. It is suggested that this selection is based on the inorganic nature of fresh ash particles, assuming initial collection for feeding: as ash particles at the sea bed develop an organic component, such selection will disappear.     

Effects of light reduction on growth of the submerged macrophyte Vallisneria americana and the community of root-associated heterotrophic bacteria

A shading experiment was conducted over a growing season to measure the effects of light reduction on Vallisneria americana in Perdido Bay on the Florida-Alabama border and to determine the response of heterotrophic bacteria in the rhizosphere. Plants subjected to 92% light reduction showed the most pronounced effects in chlorophyll a concentration, above- and below-ground biomass, and leaf dimensions. These results further suggested that the V. americana life cycle, as exhibited in temperate waters, was impaired. Heterotrophic bacteria were enumerated and identified (i) from the roots and sediments of fully illuminated plants and from unvegetated sediments at three intervals and (ii) from the roots of plants that have been subjected to 92% light reduction for 3 months. Up to two orders of magnitude greater numbers of bacteria were enumerated from root samples than sediment samples on a dry weight basis. Bacteria enumerated from the roots of plants subjected to light reduction (1.3±1.1×10⁸ CFU g⁻¹) were significantly higher than numbers of bacteria enumerated from the roots of fully illuminated plants (4.8±1.8×10⁷ g⁻¹ in the summer) or sediment samples (1.4±0.03×10⁶ g⁻¹). This suggests the roots of seagrasses stressed by light reduction provided more nutrients for bacterial growth. Higher percentages of Gram-negative bacteria were isolated from roots (up to 85% in the fall) than sediments (0-15%). Examination of isolates for traits characteristic of rhizosphere bacteria (siderophore production, formation of the phytohormone indole-3-acetic acid, and antifungal activity) did not show a clear distinction between root-associated and sediment isolates. Taxonomic identifications of root-associated bacteria based on MIDI analysis of fatty acid methyl esters were consistent with bacteria known to be associated with other plants or found at oxic-anoxic interfaces. In addition, the bacterial identifications showed most species were associated with only roots or only sediments. These results support studies suggesting seagrass rhizospheres harbor distinct bacterial communities.     

Influence of Chironomidae (Diptera) faecal pellet accumulation on lake sediment quality and larval abundance of pestiferous midge Glyptotendipes paripes

Two opposite distribution patterns of larval Glyptotendipes paripes in relation to organic carbon content in sediments of central Florida lakes were discovered. In a majority of examined lakes, G. paripes larvae were most abundant in sand sediment and their density rapidly declined with increased carbon content (type 1 lakes); however, in some cases the opposite was true (type 2 lakes). To elucidate this anomaly, field-collected organic sediments from types 1 and 2 lakes and sand sediment were studied for G. paripes development in the laboratory. Type 1 organic sediment consisted predominantly of fine particles (<0.25 mm diameter) with low dissolved oxygen levels, whereas type 2 organic sediments consisted primarily of chironomid large faecal pellet aggregates (>0.25 mm diameter), with significantly higher levels of dissolved oxygen concentrations that were similar to sand sediment. Type 2 organic sediment and sand sediment were conducive to higher survival of G. paripes larvae than fine organic sediment. The larvae in type 2 organic sediment produced longer tubes than in other sediment types. This observation indicates that accumulation of chironomid faecal pellets in lake sediments may change physical properties, such as dissolved oxygen level and consequently alter conditions for survival of chironomid larvae and possibly other benthic fauna.     

Experimental study on the influence of bioturbation performed by Nephtys caeca (Fabricius) and Nereis virens (Sars) annelidae on the distribution of dinoflagellate cysts in the sediment

Dinoflagellates include noxious microalgae responsible for the formation of toxic red tides and the poisoning of molluscs and crustaceans, resulting in important economic losses. As a consequence, the life cycle of these algae has been extensively studied, but the dormancy phase (cyst) in the sediment record is little known. In the intertidal zone, bioturbation, an important biological process resulting from the activities of benthic fauna, significantly influences the movement of particles in the sediments. Laboratory experiments have allowed comparing and quantifying the movements of fluorescent microspheres resulting from the activity of two polychaetes annelidae, Nereis virens and Nephtys caeca. The particles, which simulate 45 µm diameter dinoflagellate cysts, are deposited in flat aquaria at the surface or deep in the sediment. Photographs of the aquaria were taken at regular intervals, to observe, in a non-destructive manner, the movement of the particles and to calculate, using adapted software, the optical reworking coefficient (ORC) over time. A difference appears between the movements of the particles generated by both species of polychaetes. Nereis virens create “permanent” galleries that carry the microspheres deeply in the sediment during the digging, bioirrigation and feeding, and Nephtys caeca homogenize the particles in the first centimetres of sediment during its erratic movements. The study shows that the bioturbation generated by these organisms can modify the distribution of the 45 µm diameter dinoflagellate cysts in the sedimentary column, burying them or raising them back to the water-sediment interface.     

Evidence for the influence of seagrasses on the benthic nitrogen cycle in a coastal plain estuary near Beaufort,North Carolina (USA)

A study was undertaken to evaluate the interrelationship between the presence of seagrasses, Zostera marina and Halodule wrightii, and the physical and chemical properties of sediments in a coastal plain estuary near Beaufort, North Carolina. In sediments underlying a cover of seagrass, silt-clay, organic matter, exchangeable ammonium, ammonium dissolved in pore waters and total nitrogen were larger than in unvegetated profiles. The magnitude of the physical and chemical properties of sediments varied according to the location of the station in relation to the vegetation, as well as the continuity in the distribution of the seagrass. The largest pools of nitrogen, the finest sediment texture, and the greatest organic matter content were in sediments associated with the mid bed regions of seagrass meadows, intermediate at the edges of the bed and small isolated patches of grass, and least in unvegetated substrate.General conclusions from this study are: 1) once established, seagrasses appear capable of modifying the sediment texture as well as the organic matter and nitrogen content; 2) nitrogen accumulates beneath the vegetation suggesting that vegetated sediments are sinks; however, functional recycling mechanisms seem to be operating as suggested by the larger magnitude of remineralized nitrogen in the vegetated profiles; and 3) the establishment of seagrasses in this geographical region are not necessarily restricted by the sediment properties measured in this study. These data and conclusions are discussed in regard to an application of contemporary theories of ecosystem development to seagrass systems.Contribution Number 82-22-B     

Sediment-associated cues for larval settlement of the infaunal spionid polychaetes Polydora cornuta and Streblospio benedicti

Do patchy distribution patterns of infaunal polychaetes result from active site selection of larvae influenced by sediment-associated microbial cues? This hypothesis was tested with still-water laboratory settlement assays revealing the acceptance or rejection of polychaete larvae to qualitatively different sediments. Laboratory brood cultures of the spionid polychaetes Polydora cornuta and Streblospio benedicti yielded a sufficient number of larvae with planktotrophic development for bioassays. High settlement rates (75-95%) of test larvae were observed in response to natural sediment. Sterilization of natural sediment significantly decreased settlement of P. cornuta (25-55%) while combustion of sediment significantly decreased the settlement rate in both species (5-50%). Differences in settlement responses to sediments treated by sterilization or combustion most likely resulted from a variety of factors such as modified sediment fabric, grain size distribution and quantity of adsorbed organic matter. To experimentally address the potential role of microorganisms and microbial metabolites as mediators of larval settlement, ashed sediment was inoculated with viable microorganisms obtained from natural sediment. In both polychaete species, this treatment significantly increased larval settlement in comparison to the control of ashed sediment indicating that larval settlement was at least partially mediated by the presence of microorganisms associated with sediment.     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Distribution,substratum preference and burrowing behaviour of Lovenia elongata (Gray) (Echinoidea: Spatangoida) in the Gulf of Elat ('Aqaba), Red Sea

Lovenia elongata (Gray), the Indo-West Pacific spatangoid, was studied in the Gulf of Elat ('Aqaba) at the northern end of the Red Sea. The occurrence and local distribution of this macrobenthic burrowing species was determined and related to the particle size distribution of the substrata in which it was found. Its habitat preference is for clean, grassless, sandy bottoms in intertidal areas (beaches and sandbars), and on sublittoral level bottoms of protected bays and lagoons sheltered from heavy wave action. The preferred sediments were found to range from medium to fine sand. The population density, size frequency distribution, growth, reproduction, recruitment, and mortality of a sublittoral population of L. elongata (at Wadi Taba, Sinai) were studied.The mode of life of this burrowing heart urchin was observed in situ and in aquaria. Its burrowing and emergence behaviour are described. Experiments were carried out on the effect of particle size on burrowing behaviour. There are significant differences in the ability of L. elongata to manipulate substrata of varying grain size. The burrowing process was found to be most rapid in natural sand of medium-fine composition, corresponding to the normal sediment of the urchin's habitat. Such sediments were also found to be suitable for successful larval settlement; juveniles did not survive in coarser sand.Differences in population density and spatial distribution between various size groups are attributed to differences in their ability to manipulate the substratum. L. elongata is found to be morphologically suited to sand, particularly medium-fine sand; however, it is restricted by its morphology to shallow burrowing, since it lacks specialized mechanisms for deeper burrowing and funnel building.     

Imaging Oxygen Distribution in Marine Sediments. The Importance of Bioturbation and Sediment Heterogeneity

The influence of sediment oxygen heterogeneity, due to bioturbation, on diffusive oxygen flux was investigated. Laboratory experiments were carried out with 3 macrobenthic species presenting different bioturbation behaviour patterns: the polychaetes Nereis diversicolor and Nereis virens, both constructing ventilated galleries in the sediment column, and the gastropod Cyclope neritea, a burrowing species which does not build any structure. Oxygen two-dimensional distribution in sediments was quantified by means of the optical planar optode technique. Diffusive oxygen fluxes (mean and integrated) and a variability index were calculated on the captured oxygen images. All species increased sediment oxygen heterogeneity compared to the controls without animals. This was particularly noticeable with the polychaetes because of the construction of more or less complex burrows. Integrated diffusive oxygen flux increased with oxygen heterogeneity due to the production of interface available for solute exchanges between overlying water and sediments. This work shows that sediment heterogeneity is an important feature of the control of oxygen exchanges at the sediment-water interface.     

The functional group approach to bioturbation: The effects of biodiffusers and gallery-diffusers of the Macoma balthica community on sediment oxygen uptake

The Macoma balthica community, which is widely distributed in intertidal soft sediments bordering the north Atlantic, is dominated by two functional groups with different sediment mixing modes: the biodiffusers M. balthica and Mya arenaria and the gallery-diffuser Nereis virens. To compare the effects of these two groups on sediment oxygen uptake rates, we used experimental microcosms with identical biovolumes to measure the influence of each species on oxygen uptake. The two biodiffusers had similar effects on oxygen uptake in spite of different space occupation and different feeding, ventilation and burrowing modes. Biodiffusers and gallery-diffusers had different effects on oxygen uptake. Periodic ventilation by the gallery-diffusers stimulated the oxygen uptake by the sediment more than the steady activities of the biodiffusers. Temporal variation in oxygen fluxes in bioturbated microcosms was linked to construction and maintenance of biogenic structures. The results confirm that the functional group approach to bioturbation is a useful tool for quantifying the effects of intertidal benthic communities on benthic fluxes.     

Lower Miocene bathyal and submarine canyon ichnocoenoses from Northland, New Zealand

Thalassinidean crustacean remains ( Callianassa ) are recorded from lower Miocene burrow networks belonging to the ichnogenus Thalassinoides . They were produced at mid bathyal depths of 1000–3000 m. Mid to lower bathyal basinal sediments and inferred outer neritic to upper bathyal submarine canyon sediments contain sparse ichnocoenoses composed entirely of feeding and dwelling structures, produced by burrowing polychaetes, echinoids and possibly sipunculoids and also by bivalves escaping burial during rapid sedimentation. These two ichnocoenoses are similar to those of proximal turbidite sequences, and this is considered a response to similar rates of sedimentation, water agitation and coarseness of sediment rather than a similarity in depth. An outer neritic to upper bathyal canyon wall ichnocoenosis was developed in semi-consolidated lutites and arenites by burrowing polychaetes, decapod Crustacea (shrimps and crabs) and possibly amphipods and other organisms. Inclined, but randomly oriented Rhizocorallium occur in the canyon wall, and their presence at these depths is explained by inferred abnormally high water turbulence and abundant suspended food matter as well as a favourable semi-consolidated lutite substrate.     

Effect of fish predation on intertidal benthic fauna is modified by crab bioturbation

The burrowing crab Chasmagnathus granulatus is an important bioturbator in SW Atlantic estuaries where they generate dense and extended intertidal beds. Its bioturbation leads to profound changes in the structure, quality and dynamics of sediments with concomitant impacts on the entire benthic community. In this study, we evaluate whether the presence of this crab affects the predator-prey interaction between juvenile fishes and their benthic prey. Gut content and benthic prey selection by juvenile fishes inside and outside crab beds were evaluated, and predation effect was experimentally contrasted between areas using fish exclosures. The results show that in crab beds the percentage of fish with empty guts was lower and the number of polychaetes consumed by fish higher than outside crab beds. The silverside Odontesthes argentinensis and the catfish Pimelodella laticeps fed on larger polychaetes outside than inside crab bed areas, while the white mouth croaker Micropogonias furnieri preyed upon larger polychaetes inside crab beds. In addition, field experiments shows that fish predation decreases polychaete abundances only in crab beds. These results suggest that crab bioturbation facilitate fish predation on benthic prey.     

Composition, abundance and stratification of soft-bottom macrobenthos from selected areas of the Ross Sea shelf (Antarctica)

Composition, abundance and stratification of soft-bottom macrobenthos were studied at three selected sites on the Ross Sea shelf (Antarctica) with different geomorphology and sedimentation regimes. Sites A (southwest Ross Sea, 810 m depth) and B (Joides basin, 580 m depth) were characterized by biogenic mud and clay sediments, whereas site C (Mawson bank, 450 m depth) featured sandy sediments mixed with a conspicuous biogenic component characterized by shells and tests of calcareous invertebrates (mainly barnacles of the genus Bathylasma). The macrofauna of sites A and B was mainly composed of infaunal polychaetes and bivalves. The assemblages comprised both surface and sub-surface deposit feeders, including some conveyor-belt polychaetes (Maldanidae and Capitellidae) that are responsible for high sediment mixing and bioturbation. The macrobenthos of site C was dominated by crustaceans, polychaetes and echinoderms (ophiuroids), and mainly by filter feeders and epifaunal or interstitial forms. Abundances were higher (up to 1040 ind. m⁻²) at site B than at sites A and C (430 and 516 ind. m⁻², respectively). At sites A and B the benthos was mainly concentrated in the upper 5 cm of the sediment, and abundances declined sharply in the deeper sediment layers. These results indicate a high degree of consistency between sediment features and benthic community structure, which are both strongly related to local hydrography and bottom dynamics. Sites A and B represent areas where the organic input to the seafloor by vertical sedimentation from the upper water column is high. Site C, however, is characterized by high sediment instability and food particles advecting mainly horizontally. The community is more physically controlled by unpredictable, and probably frequent, disturbance events (e.g., bottom turbid currents, sediment reworking and displacement). Individuals were relatively small, indicating that probably they are not able to grow up to the adult size and reproduce. The community may be represented by “pseudopopulations” depending on the settlement of larvae invading from neighbouring areas. Accepted: 23 October 1998