Methods for monitoring tidal flushing in large animal burrows in tropical mangrove swamps

The typically anaerobic nature of mangrove sediments provides significant challenges to the mangrove trees and biota inhabiting them. The burrowing activities and flow of water through the numerous and complex animal burrows perforating the sediments of mangroves have a major influence on the biogeochemistry of the sediments and are important to the enhancement of nutrient and oxygen exchange. Two new methods are presented for monitoring the tidal flushing of Sesarma messa and Alpheus cf macklay burrows in a Rhizophora stylosa mangrove forest – by measuring oxygen content of burrow water and by determining the change in fluorescence of a dye tracer through tidal inundation. A case study using the first of these showed oxygen consumption rates at the burrow wall deep within the burrow were found to be between 210 and 460 μmol O₂ m⁻² h⁻¹. The influx of oxygen during a flood tide was found to be significant and indicated that approximately 40% of the burrow water is flushed during a single tidal event. However, the high consumption rate of oxygen within the burrow resulted in the oxygen concentration remaining at or below one-third of the oxygen content of the flooding tidal water. A test application of the second method, using rhodamine dye as a tracer, indicated that the exchange of water between the burrow and the flooding tide was found to be in the order of 30% of the burrow volume. These new techniques provide a means to further study the nutrient exchange within these burrow systems and verify the initial findings that several tidal inundations are necessary to completely flush the burrows.     

Distribution patterns of the ghost crab Ocypode cursor on sandy beaches of a tropical island in the Cabo Verde archipelago,Eastern Central Atlantic

The spatial distribution of the ghost crab Ocypode cursor was determined for beaches on eastern Boa Vista Island, Cabo Verde Archipelago. The main objectives were to analyse the across-shore distribution by means of burrow counts and to identify preferential zones and spatial segregation. Six beaches were investigated using 20 across-shore beach transects. It was demonstrated that ghost crabs occur throughout the entire intertidal zone, with higher densities in a 9–13 m band above the waterline. There was an exponential relationship between the carapace length of captured crabs and the diameter of their burrows. Mean burrow diameter was correlated with burrow density and increased linearly with distance from the waterline, although small burrows, representing individuals smaller than the median size, occurred across the beach. Burrow diameter and depth were positively correlated, with larger holes indicating larger crabs and deeper burrows. Deeper burrows were more common in the upper intertidal zone. These results are similar to those described for ghost crab populations elsewhere and contribute to an understanding of the ecology of the species in the local context.     

Intertidal mangrove boundary zones as nursery grounds for the mud crab Scylla serrata

The demand for mud crab Scylla serrata (Forsskål 1775) in the global market has increased, hence there is growing momentum to farm the species in Africa. Aquaculture production in Kenya and elsewhere in East Africa currently relies on juvenile seeds sourced from the wild. Wild-seed collection calls for management of the juvenile crab industry founded on knowledge of the species’ ecology, so as to achieve a sustainable seed supply and recruitment to the capture fishery. This study investigated the tidal, diurnal and seasonal occurrence of juvenile crabs in three habitats (intertidal-flat boundary zones, inside the mangroves, and in channels) in small creeks (Mida, Kilifi and Mtwapa) and Gazi Bay, on the coast of Kenya. Sampling was done with scoop nets and seining at receding tides and via burrow searches at low spring tides (day and night). Juveniles in the mangrove/intertidal-flat boundary zone were found sheltered under mangrove leaves or debris, or in shallow burrows during low spring tides, whereas at receding tides they could be seen moving out with the tide or searching for sheltering substrate or burrows. Catch per unit effort at high-abundance sites varied between 59 and 68 crabs fisher^?1 day^?1. More juvenile crabs, sized 10–80 mm internal carapace width (ICW), occurred at night during the receding tide. Net-seining was effective in the collection of juvenile mud crabs <30 mm ICW, whereas burrow sampling was effective for gathering larger crabs. However, collection of juveniles by seining reduced the quality of the crabs caught due to frequent loss of chelipeds, as compared with retrieving individual crabs by searching burrows. Some juveniles collected in the intertidal-flat boundary-zone habitat were either in the process of moulting or had just moulted, indicating the significance of this habitat for mud crabs at this physically delicate life stage.     

The tidal flushing of multiple-loop animal burrows

Tidal flushing of animal burrows in mangrove swamps provides an effective and important mechanism for transport of salt and other soluble substances. The burrows have complex morphologies consisting of multiple loops. Using established computational modelling techniques, burrow geometries were simulated from characteristic burrow dimensions and the effect of multiple loops on flushing is studied. The computational models show that flushing is enhanced in multiple-loop burrows as upper loops can be completely evacuated, increasing the volume of burrow water removed. The models indicate that there may be a depth limit to which flushing occurs in complex burrow structures. Periscope loops extend further than one characteristic loop depth and are shown to affect the flushing of burrows if the surface-water plume penetrates to their lower depth. Periscope loops with surface openings on the downstream side of the burrow, relative to the tidal inundation, most frequently experience this condition and thus have a great impact on burrow flushing. Field measurements of salinity agree with the hypothesis that there is a depth limit of flushing for complex burrows that is independent of burrow salinity, and suggest a value of 30–35 cm. The measurements also support the suggestion that significant flushing of burrows occurs within a single tidal event. An experiment considered the movement of animals across the interface created by low-salinity surface water and high-salinity burrow water after a partial flushing event. The motion enhances the effective diffusion coefficient of salt by a factor of at least 10², as compared with free diffusion of salt in water, resulting in a more even distribution of dissolved salt in the burrow water between tidal inundations. This distribution may lead to an increase in the depth to which flushing occurs during subsequent inundations.     

The Southwest Atlantic intertidal burrowing crab Neohelice granulata modifies nutrient loads of phreatic waters entering coastal area

Along the coastal areas of the Southwest Atlantic estuaries and embayments, phreatic water often circulates through very extended areas (up to several hundred meters perpendicular to the coast), dominated by dense assemblages of deep burrows of the crab Neohelice granulata (formerly Chasmagnathus granulatus). This crab inhabits the intertidal area, from mudflats to marshes vegetated by species of Spartina, Sarcocornia and Juncus, generating extensive burrowing beds where burrow density may reach up to 60 burrows m⁻². Since the lower limit of the crab burrows is usually the water table, we investigated through field experiments the effect of N. granulata and their burrows on the chemical characteristics of this phreatic water. Water analysis from experimental (1) occupied burrows (with crabs), (2) unoccupied burrows (where crabs were excluded), and (3) sediment pore water show remarkable differences. Water oxygenation, and nitrate, ammonium and sulphate concentrations inside occupied burrows were higher than in the water inside unoccupied burrows or pore waters. Moreover, directed sampling of phreatic water entering and leaving the crab bed, shows that dissolved inorganic nitrogen concentration is enhanced as the water crosses the crab bed. These results may be ascribed to the fact that in the salt marsh the crabs spend most of their time within burrows, where presumably they store food (plants) and defecate. These activities generate an area of accumulation of excrements and nutrients in different decomposition states. The present work shows a novel way by which bioturbating organisms can affect nutrients exportation from salt marshes to the open waters.     

Novel mechanism of stream formation in coastal wetlands by crab–fish–groundwater interaction

A novel mechanism of stream development in coastal wetlands was observed in the form of interaction between the burrowing crab (Chasmagnathus granulata) and groundwater seepage. This process may eventually be further promoted by the predatory action of the white croaker fish (Micropogon opercularis). These so-called crab streams occur on the banks of major tidal channels in a salt marsh of the Bahía Blanca Estuary (Argentina). As the tide recedes, water is retained in the crab burrows on the marsh surface. The presence of lateral burrows produced by crab recruits promotes subterranean piping flow driven by the hydraulic head in the vertical burrow. Groundwater generally seeps along the tidal channel banks but is particularly prominent and concentrated where crab burrows perforate the banks. Where predatory attacks by the croaker fish enlarge the burrow mouths, the drainage rills become wider and deeper because more seeping water is collected in the shallow crater from where it is funnelled into the developing streams. The crab stream is then produced by water draining from the crab burrow mouth, acting initially by cut-back erosion and followed by forward erosion of the relatively unconsolidated muddy sediments.     

Burrow Structure and Burrowing Activity of the Funnel-feeding Enteropneust Balanoglossus aurantiacus in Bogue Sound, North Carolina, USA

Abstract. This study of the enteropneust, Balanoglossus aurantiacus , in Bogue Sound, NC, provides information on the general structure and variability in geometry of its burrow, B. aurantiacus employs a funnel at the anterior end of its burrow to cave in surface sediments. It ingests, processes, then deposits these sediments back on the surface as fecal castings. Measurements made on resin casts of burrows and on photographs of surface features (burrow openings and funnels) revealed a new and consistent pattern of burrow construction. This pattern distinguishes B. aurantiacus burrows from those reported for other funnel feeders and enteropneusts (except, perhaps, for B. clavigerus). Laboratory and field observations revealed that, within its burrow, B. aurantiacus is elongated, can double-back on itself, and can use anterior burrow openings for subsequent fecal castings. Tests for significant changes in location of surface features gave estimates of how frequently individuals in aquaria and field populations abandon old burrow locations and construct new burrows. Field observations showed that B. aurantiacus will reopen a blocked burrow opening.     

Spilled Oil and infaunal activity—modification of burrowing behavior and redistribution of oil

A series of experiments in Willapa Bay, Washington, indicates the degree to which the presence of spilled oil modifies the burrowing behavior of infauna and the extent to which the animals redistribute oil into interridal sediment. Small amounts of North Slope crude oil introduced at low tide directly into burrow openings (mostly made by the crustacean Callianassa) resulted in a limited and temporary reduction in the number of burrow openings. In contrast, a layerof oil-saturated sand 1 cm thick buried about 5 cm below the sediment surface sharply reduced the number of burrow openings. After a year, the few new burrows penetrated only the margins of the experimental plot, and bioturbation below the buried oil saturated sand layer declined dramatically.The experiments suggest that small amounts of oil temporarily stranded by tides in themselves have no long-range effect on burrowing behavior. The fauna, however, are capable of introducing measurable amounts of oil into the subsurface, where it is retained long after the rest of the stranded oil has washed away. A buried layer of oil-saturated sand greatly reduces infaunal activity; the oil presents an effective barrier that can persist for years.The oil incorporated into the sediment from burrow openings showed evidence of degradation after 7 months. In contrast, the layer of buried oil remained essentially undegraded after a period of two years, even though oil in lower concentrations above the layer was degraded after a period of one year. This variation in degree of dgradation of the buried oil, as well as the heterogeneity of oil distribution wherever the oil has been incorporated from the surface, emphasises the importance of careful sampling in any attempt to locate or monitor the presence of spilled oil in the substrate.     

米草属植物Spartina angilica和Spartina alterniflora引种后江苏海岸湿地生态演化的初步探讨

选取江苏王港潮滩的盐蒿滩、大米草滩及互花米草滩,分析这三类滩面的地面以上生物量、初级生产力、动物洞穴参数和沉积物粒度,探讨大米草和互花米草引种区的湿地生态系统相对于盐蒿滩的变化.分析结果表明,互花米草滩的生物量远大于盐蒿滩和大米草滩;单位面积上动物洞穴数量相近,但洞穴大小有一定差异,互花米草滩动物洞穴稍大,数量也相对较多,这可能与初级生产力的提高有关.表层底质的粒径以互花米草滩为最细,盐蒿滩最粗,这种分布状况与互花米草引种前不同,说明互花米草促进了细颗粒物质的堆积.互花米草的引种在江苏海岸具有促淤和提高初级生产力的作用,而它对湿地生态系统的结构和功能的影响需要进一步研究.     

Observations on the Burrows and Burrowing Behaviour of the Red Band-Fish, Cepola rubescens L.

Abstract. The red band-fish, Cepola rubescens L., lives in burrows in sublittoral muddy sediments. The authors first presented information on the burrows of this species in the 1970s. The present paper presents new information on burrow structure, describes the method of excavation, and comments on the bioturbatory significance of the species. The work derives from field and laboratory studies. A burrow typically consists of a vertical shaft which opens into an expanded terminal chamber. In some cases a side shaft may be added. The paper includes a size analysis of 130 burrows measured in the field and detailed morphological information from a selection of burrows which were cast with polyester resin. The fish burrows are frequently intersected by the burrows of other species and interspecific associations may develop. Burrow size reflects the size of the occupant and may approach 1 m in depth. The biogenic movement of water and particles to this depth is often overlooked in bioturbation studies and is discussed. Burrow distribution is aggregate, which has implications for the bioturbatory impact of the species. Burrows are constructed by mouth excavation and this is described in detail. Fish transport fine material within their mouths and coarse material is grasped in the jaws. Large spoil heaps occur at burrow openings. One obvious effect of this bioturbatory activity at the field site was the redistribution of coarse material (shell gravel) from depth to the sediment surface.     

Reproductive ecology of the mudskipper Boleophthalmus pectinirostris

The reproductive ecology of the mudskipper Boleophthalmus pectinirostris was investigated during the spawning season in the Funing Bay, Fujian, China. The fish burrows were basically Y-shaped and had two openings onto the mudflats. Part of the intersection at the center of the burrow was dilated to form a "spawning chamber". The dissolved oxygen concentrations (0.40~0.65 mg/dm3)of water 15 cm deep inside the burrows were much lower than those (5.96~6.19 mg/dm3) of intertidal pool water. Water temperatures inside the burrows were much lower than those of intertidal pools. Field investigations indicated that eggs were laid on the inner wall and ceiling of the spawning chamber by means of filamentous attachments, and a male stayed inside the burrows to guard it. Field investigations also suggested that fish of both sexes constructed mud burrows by themselves but that the spawning chamber was made only in the male burrow. The male attracted a female to his burrow for mating and spawning. There was no water in the spawning chamber, and thus the eggs were exposed to the air in the chamber. Changes in spawning readiness and gonadosomatic index indicated that synchronization of spawning was related to the semi-lunar periodicity, and this is the first report of this relationship in the mudskippers.     

Observations on the Burrows and Burrowing Behaviour of: Squilla mantis L. Crustacea: Stomatopoda

Abstract. The burrows of Squilla mantis are described from polyester resin casts taken at two Adriatic sites near Ancona. Typically, burrows had two openings, one of which was larger than the other, and were basically U-shaped. An interesting association with the little-known caridean shrimp Athanas amazone was observed. The sediments occupied by S. mantis were silty sands and sandy muds. Field observations of S. mantis and other species on the same grounds are presented and some information on burrow density is given, derived from burrow counts using underwater television. Burrowing behaviour is described from observations of animals transferred to laboratory aquaria.     

Sediment dynamics modulated by burrowing crab activities in contrasting SW Atlantic intertidal habitats

Biogenic bottom features, animal burrows and biological activities interact with the hydrodynamics of the sediment–water interface to produce altered patterns of sediment erosion, transport and deposition which have consequences for large-scale geomorphologic features. It has been suggested that depending on the hydrodynamic status of the habitat, the biological activity on the bottom may have a variety of effects. In some cases, different bioturbation activities by the same organism can result in different consequences. The burrowing crab Neohelice granulata is the most important bioturbator at SW Atlantic saltmarshes and tidal plains. Because of the great variety of habitats that this species may inhabit, it is possible to compare its bioturbation effects between zones dominated by different hydrodynamic conditions. Internal marsh microhabitats, tidal creeks bottoms and basins, and open mudflats were selected as contrasting zones for the comparison on a large saltmarsh at Bahía Blanca Estuary (Argentina). Crab burrows act as passive traps of sediment in all zones, because their entrances remain open during inundation periods at high tide. Mounds are generated when crabs remove sediments from the burrows to the surface and become distinctive features in all the zones. Two different mechanisms of sediment transport utilizing mounds as sediment sources were registered. In the first one, parts of fresh mound sediments were transported when exposed to water flow during flooding and ebbing tide, with higher mound erosion where currents were higher as compared to internal marsh habitats and open mudflats. In the second mechanism, mounds exposed to atmospheric influence during low tide became desiccated and cracked forming ellipsoidal blocks, which were then transported by currents in zones of intense water flow in the saltmarsh edge. Sedimentary dynamics varied between zones; crabs were promoting trapping of sediments in the internal saltmarsh (380 g m⁻² day⁻¹) and open mudflats (1.2 kg m⁻² day⁻¹), but were enhancing sediment removal in the saltmarsh edge (between 10 and 500 g m⁻² day⁻¹ in summer). The implication is that biologically mediated sedimentological changes could be different among microhabitats, potentially leading to contrasting geomorphologic effects within a particular ecosystem.     

Seagrass as the main food source of Neaxius acanthus (Thalassinidea: Strahlaxiidae), its burrow associates, and of Corallianassa coutierei (Thalassinidea: Callianassidae)

Burrows of the thalassinidean shrimps Neaxius acanthus and Corallianassa coutierei are striking aspects in tropical seagrass beds of the Spermonde Archipelago, Indonesia. Burrow construction, behaviour, burrow type and associated commensal community were investigated to clarify the ecological role and food requirements of these shrimps and their commensals. Gut content analysis and stable-isotope data were used to unravel the food sources and the trophic interactions among the commensal community.Individuals of Neaxius acanthus were caught on Bone Batang Island. In narrow aquaria filled with sediment they constructed burrows resembling those found in the field. During burrow construction and maintenance only little sediment was brought to the surface, most was sorted and compacted to create a distinct lining. Maintenance work by single shrimps typically took about 5 min, after which the shrimp walked up to the entrance and rested for a similar period of time. There were no differences in behaviour between day and night. Intrasexual encounters inside the burrow were characterised by a high level of aggression and all resulted in one participant being driven out of the burrow. Intersexual encounters led to coexistence with both animals taking turns in burrow maintenance and guarding the entrance. Offered seagrass leaves were pulled underground, cut into pieces and eventually integrated into the lining. Burrows of Corallianassa coutierei resembled a deep U-shape. Chambers branching off halfway down and at the deepest point contained seagrass fragments. All steep parts of the burrow were lined similar to burrows of N. acanthus.No commensals were found associated with Corallianassa coutierei. However, burrows of Neaxius acanthus in the field typically contained a pair of shrimps, up to 8 individuals of the commensal bivalve Barrimysia cumingii and large numbers of gammarid amphipods. Other animals found associated with the burrow were the goby Austrolethops wardi, a palaemonid shrimp species and two species of tube-building polychaetes, one of which was also found as an epibiont on N. acanthus.Stable-isotope and gut content analyses indicate that the diet of Neaxius acanthus, its commensal Austrolethops wardi, and Corallianassa coutierei is mainly derived from detrital seagrass leaves, with a potential contribution of sediment organic matter and seagrass epiphytes. In contrast the isotopic signature of Barrimysia cumingii suggests the presence of symbiotic sulphide metabolism bacteria. This study underlines that, besides their interactions with the surrounding ecosystem, thalassinid shrimp burrows play an important role as a sub-habitat with a unique associated fauna.     

Habitat-related differences in the pre-copulatory guarding and copulation behavior of Neohelice granulata (Brachyura,Grapsoidea, Varunidae)

Mating behavior strategies of brachyuran crabs are flexible depending on the habitat and social context. We evaluate pre-copulatory guarding in Neohelice granulata and describe the unusual copulatory behavior. Laboratory experiments were conducted to test whether factors such as male size, presence of burrows, a male-biased operational sex ratio (OSR) and female mating history affect the copulation duration in two different study areas. Females initiated a searching behavior approximately 20–26 h before they became receptive by assessing large male burrows and displaying a courtship behavior (“flirting”) in its entrance. Once the female entered inside the male burrow, pre-copulatory guarding was observed until the female became receptive and successful copulation occurred immediately after. This species shows some very unusual copulatory behavior involving eyestalk grasping not seen in any other varunid crab. The copulation duration was independent of male size and the presence of burrows, although it was dependent on the study area, the OSR and the female mating history. Thus, copulation duration is a flexible character in this species which varies with habitat and biological factors such as mating history and sex ratio.     

Variation in the post‐copulatory guarding behavior of Neohelice granulata (Brachyura,Grapsoidea, Varunidae) in two different habitats

Crab guarding behaviors are influenced by the biological and physical characteristics that each population experiences in a particular habitat. We studied post‐copulatory guarding behavior and the factors affecting it, such as the presence of burrows, female receptivity duration, operational sex ratio (OSR) and male size, in two different populations of Neohelice granulata: Mar Chiquita coastal lagoon (MCL) and San Antonio Oeste (SAO). Laboratory experiments showed that post‐copulatory guarding duration varied between the study areas. In MCL, it depended upon the presence of burrows and female receptivity duration, whereas it was independent of the OSR and male size. In SAO, it was dependent upon the OSR and female receptivity duration, whereas it was independent of the presence of burrows and male size. Thus, N. granulata males display different reproductive strategies related to their post‐copulatory guarding behavior pattern in different habitats. These strategies are related to the fact that this species constructs and defends a burrow that provides shelter and a place to mate, something that other varunids do not have.     

Infaunal burrow ventilation and pore-water transport in muddy sediments

The ventilation of burrows by tube-dwelling benthos is understood to be important in determining rates of exchange of solutes between the sediment and overlying water. However, few models have attempted to link the burrow ventilation behavior of tube-dwelling organisms with their geochemical consequences. The classic cylinder model of bioirrigation in muddy sediments (Aller, R.C., 1980. Quantifying solute distributions in the bioturbated zone of marine sediments by defining an average microenvironment. Geochimica et Cosmochimica Acta 44, 1955–1965) links pore-water processes and burrow sizes and distributions in the sediment by assuming that burrows are fully flushed. The equivalence between the cylinder model and the more commonly used one-dimensional non-local exchange model depends upon this assumption. However, this assumption has seldom been tested in the field. We have extended the cylinder model of bioirrigation to include burrow ventilation activities of organisms. Burrow ventilation is modeled as a simple non-local exchange of burrow water with overlying water. Model simulations indicate that burrow ventilation has a large effect on vertical profiles and fluxes of solute tracers. We collected data on burrow geometry in the field by CT-scanning freshly collected sediment cores. At the same study sites, we measured activity profiles of ²²²Rn, a naturally occurring radionuclide tracer of pore-water transport. With model geometry independently constrained, we tested the model by comparing our model-predicted profiles with measured profiles. Our results demonstrate that burrows in the field are not fully flushed. Our estimated burrow ventilation rates compare favorably with previous laboratory measurements. The inclusion of realistic burrow ventilation in this pore-water transport model strongly affects modeled solute profiles and fluxes. We demonstrate how model parameters can be determined from field samples and present a model that more realistically simulates pore-water transport processes in muddy sediments.     

The sediment-water interface increment due to the complex burrows of macrofauna in a tidal flat

The architecture of macrofaunal burrows and the total area of the sediment-water interface created by biogenic structure were investigated in the Donggeomdo tidal flat on the west coast of Korea. Resin casting methods were applied to recover burrows of four dominant species, Macrophthalmus japonicus, Cleistostoma dilatatum, Perinereis aibuhitensis, and Periserrula leucophryna, and whole burrows within the casting area at three sites in different tidal levels.P. leucophryna excavated the largest burrow in terms of a surface area among them. In the case of whole burrow casting, the space occupied by the biogenic structure was extended into deeper and expanded more greatly at the higher tidal level. In the uppermost flat, the burrow wall surface area within sediment was more extensive than the sediment surface area. Increased oxygen supply through the extended interface could enhance the degradation rates of organic carbon and also change the pathways of degradation. Quantifying the relationship between the extended interface and mineralization rate and pathway requires more extensive study.     

Vehicles versus conservation of invertebrates on sandy beaches: mortalities inflicted by off-road vehicles on ghost crabs

Sandy beaches face increasing anthropogenic pressures, with vehicle traffic being ecologically highly harmful. Ghost crabs (Fam. Ocypodidae) are conspicuous on many beaches, and they have been used as a bio‐monitoring tool to measure the ecological responses to human disturbance. However, the mechanisms causing declines in crab numbers are unknown, yet conservation must target the actual impact mechanisms. Therefore, we quantified the magnitude and mechanisms of off‐road vehicle (ORV) impacts on ghost crabs, addressing three key questions: (i) Does abundance of ghost crabs respond to traffic intensity?; (ii) Can burrows protect crabs from vehicles? and (iii) Can mortalities caused by vehicles contribute to population declines? ORV‐impacts were measured on North Stradbroke Island (Australia) for Ocypode cordimanus and Ocypode ceratophthalma. Crab densities were significantly lower in areas subjected to heavy beach traffic, suggesting direct crushing by vehicles. Burrows only partially protect crabs against cars: all individuals buried shallow (5 cm) are killed by 10 vehicle passes. Mortality declines with depth of burrows, but remains considerable (10–30% killed) at 20 cm and only those crabs buried at least 30 cm are not killed by ORVs: these ‘deep‐living’ crabs represent about half of the population. After crabs emerge at dusk they are killed in large numbers on the beach surface. A single vehicle can crush up to 0.75% of the intertidal population. While conservation measures should primarily regulate night traffic, our results also emphasise that the fossorial life habits of sandy beach animals cannot off‐set the impacts caused by ORVs.     

Bioturbation effects of Corophium volutator: Importance of density and behavioural activity

Bioturbation is one of the major processes influencing ecosystem functioning. Population parameters such as species density, burrow density and species-specific life modes, determine the impact of bioturbation on the ecosystem. A laboratory experiment was developed, using microcosms mimicking a marine intertidal sediment–water interface which allowed for quantification of different population parameters. The vertical redistribution, bioturbation rate and maximum penetration depth of two sizes (41 and 129 μm) of luminophores were measured in five treatments (control, low density of burrows with and without Corophium (1989 ind./m²), and high density of burrows with and without Corophium (14,921 ind./m²)) after 1, 7 and 14 days. Results indicate that the behavioural activities of Corophium are of the utmost importance in sediment reworking, since they contributed to a five-fold increase in bioturbation rate compared to the passive transport induced by the static structure of the burrows. Furthermore, density is an important parameter because only high densities play a prominent role in particle transport and hence in organic matter processing, while the role of low Corophium densities is limited in sediment reworking. No evidence for differentiation in sediment size fractions was observed. Finally, bioturbation rates in this study were low compared to other studies, and these results suggest an influence of the tidal rhythmicity in the behavioural activity of Corophium on the bioturbation rate.