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.     

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.     

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.     

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.     

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.     

Oxygen penetration through invertebrate burrow walls in Korean tidal flat

Macrofaunal burrows increase the surface area of the sediment-water interface, which has a great impact on the metabolism of aerobic microbes and the oscillation of reduced metabolites within the sediment. Given the importance of macrofauna in surficial sediments, the aim of this study is to examine the effects of burrow architectures on dissolved oxygen diffusion rates in comparison with unburrowed sediment, and thereby to evaluate the theoretical assumption used for modeling solute distribution in the burrow system using field samples. Employing microsensors, horizontal oxygen profiles were measured on a micrometer scale around burrows of seven invertebrates in tidal flats of the west coast of Korea. Oxygen diffusion distance through the burrow walls of seven invertebrates showed spatio-temporal variation with a range of 0.6 to 2.9 mm. Two groups of burrows were identified based on their oxygen diffusive properties relative to unburrowed sediments: 1) oxygen penetration similar to that of ambient sediments and 2) clearly enhanced oxygen penetration. Differences in the diffusive properties of the burrow wall were related to the burrow depth and diameter, existence of mucus lining on the wall, sediment grain size, and tidal phases. Also inhabitant activity was an important factor affecting oxygen penetration, which is discussed in the paper. These results further demonstrate that simplified assumptions (i.e. burrow structures are viewed as direct biogeochemical extensions of the sediment-water interface) may not be exact representation of the nature.     

A deep burrowing sipunculan of ecological and geochemical importance

In late 1980s, a dense network of deep capillary burrows was reported on the Vøring Plateau, Nordic Seas, and associated with a sipunculan belonging to the genus Nephasoma. This sipunculan was responsible for rapid transport of organic matter from the sediment surface down a deep burrow network. Over 460 specimens belonging to the genus Nephasoma were collected from the deep Nordic Seas during four cruises from 2000–2005 and four species identified: Nephasoma abyssorum abyssorum, N. diaphanes diaphanes, N. diaphanes corrugatum and N. lilljeborgi. The species responsible for the deep burrows and rapid subduction of organic matter can now be confirmed as Nephasoma lilljeborgi. Deep burrows associated with N. lilljeborgi were observed on the Vøring Plateau, Bear Island Fan, Svalbard Margin and Yermak Plateau and may be a seabed feature endemic to the Nordic Seas region. N. lilljeborgi could have a significant role in influencing the ecology and geochemistry of the Nordic Seas region. It is recommended that future benthic community studies in the Nordic Seas region confirm the species identity of sipunculan specimens in order to determine the ecological and geochemical importance of the specimens.     

Burrowing and burying by the crab Macrophthalmus hirtipes

Macrophthalmus hirtipes (Jacquinot, 1853) excavates burrows under water in soft, muddy sand that remains waterlogged at low tide. The crabs are immersed for at least 4 h in every tide. The burrows are usually single, blind‐ending, slightly curved passages; their openings do not appear to be deliberately closed by the crabs. Excavation is effected by using one set of walking legs as a scoop while the other set provides sideways traction.

Burrow openings are readily obscured by water movement, so that apparent burrow density does not give a true indication of the number of crabs in an area, but only indicates how many have been active recently; the true density may be found by sieving.

The burrowing activity of M. hirtipes is compared with that of other ocypodid crabs, and the functions of burrows are discussed. A distinction is made between excavating a burrow and rapid burying, because the two behaviours are very different, and must involve different adaptations and selection pressures.     

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.     

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.     

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.     

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.     

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.     

Evidence of bioturbation in the Cap-Ferret Canyon in the deep northeastern Atlantic

In the sedimentary column, a combined quantification of burrows and macrobenthic community provides evidence of bioturbation features in the submarine canyon of Cap-Ferret between 2000 and 3000 m depth. An image-processing technique allows accurate quantification of burrow volumes with depth in the sedimentary column. The major bioturbation mode seems to be different in the channel compared to the interfluve. Macrobenthic activity is more inclined to mix the sediment in the channel in response to increased organic matter supplies. Sediment mixing leads to burrow destruction in the upper mixed layer of sediment in the canyon. Burrows are better preserved on the interfluve where mixing is slower. Under the mixed zone, the volume of recorded burrows is higher when sedimentation rate increases, as in the upper canyon. In this transition layer, the burrow volume is estimated to be between 3 and 64% of the total sediment volume depending on the sediment depth. The fill-down of numerous burrows with surface sediment by bioregeneration suggests that anaerobic degradation of fresh organic matter is dominant in this canyon. In the sedimentary column, the negative relationship between carbonate content and macrobenthic abundance confirms that carbonate dissolution is largely influenced by bioturbation.     

Pumping rates of the mud shrimpCallianassa japonica

Pumping rate of a mud shrimp,Callianassa japonica, in its burrow was measured by continuous monitoring of dye concentration in the burrow water. Measurement of dilution in two directions from stained overlying seawater to normal burrow water andvice versa, gave no significant difference in results. The rate of exchange (v) of burrow water was estimated from,v=(u _t−u ₀)V/(m−u _t−1)t, whereV is volume of burrow water,u ₀,u _t−1 andu _tis dye concentration of burrow water at time 0,t−1 andt, respectively, andm is dye concentration of overlying water. The pumping rate ranged from 0.63 to 5.46 ml min⁻¹, which corresponded to a turnover time for the burrow water of 7–51 min. Short term changes in the pumping rate were correlated to intermittent behaviour of the shrimp in the burrow.     

The distribution of zinc,nickel, manganese,cadmium, copper,and iron in some surface waters from the world ocean

Determinations of Zn, Ni, Mn, Cd, Cu, and Fe have been carried out on 51 near-shore and 38 open-ocean surface seawaters from various regions of the World Ocean.The concentrations of the trace metals have been established in the open-ocean waters, and have been used as “baselines” to evaluate trace-metal enhancement in near-shore regions. The factors by which the trace metals are enriched in near-shore regions vary from one element to another and, according to the highest concentration factors found, decrease in the order Zn = Mn > Cd = Cu = Ni.These elements exhibit differences in their distributions between near-shore and open-ocean waters, and they have been divided into two types on this basis: Type I, in which the largest number of samples in both shelf and open-ocean waters lie in the same concentration range. Zinc, cadmium, and copper are Type I elements. Type 2, in which the largest number of samples of near-shore waters lie in a higher concentration range than the largest number of samples of open-ocean waters. Nickel and manganese are Type 2 elements.The concentrations of Mn, Cd and Zn are similar in open-ocean surface waters from the South Atlantic and Indian Ocean, but Cu and Ni have higher concentrations in the former ocean.There is considerable variation in the concentrations of the trace metals in near-shore surface waters from various regions of the World Ocean. These variations are discussed in detail.     

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.     

Automatic image analysis of X-ray radiographs: a new method for ichnofabric evaluation

Automated ichnofabric analysis was performed on X-ray radiographs from five gravity cores retrieved from Late Quaternary sediments from the SW Portuguese continental slope. The image analysis program DIAna was used to detect and quantify the part of the ichnofabric consisting of pyritized microburrows, presumably Trichichnus and “Mycellia”. Abundance and orientation of the burrows were determined at 5-cm intervals. The results agree with manual counts of burrows in the coarse fraction of the sediment, and they indicate that the abundance of the pyritized burrows depends on enhanced organic carbon content, fine grain size, and low bottom water oxygenation. In contrast, no clear correlation between burrow orientation and environmental conditions was observed.     

First account of apochlorotic diatoms from intertidal sand of a south Florida beach

During the period January to December 2004, monthly intertidal sand samples were collected from John U. Lloyd State Park beach, Florida. This study is the first to report on spatial and temporal distributions of heterotrophic (apochlorotic) diatoms in a sub-tropical beach, or indeed any sandy beach. Two non-pigmented, apochlorotic diatom morphotypes (strains III and IV) were consistently isolated from two intertidal beach sites and neither conformed to the morphologies of described species. Apochlorotic diatoms were found throughout the year and formed significant peaks in April, attaining total diatom counts of 71.1 and 94.9 cells g⁻¹ wet sand, for the upper tidal zone and lower tidal zone, respectively. Valve morphology is described in detail for the two diatom morphotypes isolated from the beach sites (strains III and IV). Growth responses to changes in salinity (reflecting those of the intertidal zone) were examined in the laboratory for these diatoms. Both morphotypes were euryhaline, tolerating salinities between 10 and 50, and maximum growth rates were achieved at salinities between 20 and 40. Diatoms grew equally well whether grown in the light or the dark and showed no evidence of pigments when viewed by epifluorescence microscopy. The ability of diatoms to burrow into agar and carrageenan at different concentrations was tested. Both morphotypes were observed to develop burrows through non-nutrient agar up to twice the standard concentration (30 g agar l⁻¹). The ease with which these heterotrophic diatoms formed burrows suggests that they were digesting these complex carbohydrates (i.e. phycocolloids) as a nutrient source. Strain IV, however, failed to burrow in a different phycocolloid, carrageenan. Similar degradative actions in the field might have implications for the breakdown of recalcitrant materials or allow diatoms to effectively compete with bacteria for carbon and nutrients.     

Effect of environmental conditions on variation in the sediment-water interface created by complex macrofaunal burrows on a tidal flat

We quantified the increase in the sediment-water interface created by the burrowing activities of the resident macrofaunal community and its variation with respect to the physical conditions of the habitat on a tidal fat. We investigated environmental factors and dimensions of macrofaunal burrows with respect to tidal height and vegetation during spring and summer at three sites. A resin-casting method was used to quantify the dimensions of all burrows at each site. The dimensions of macrofaunal burrows varied both temporally and spatially and the increase in the sediment-water interface reached a maximum of 311%, ranging from 20 to 255% under different habitat conditions. The sediment-water interface depended on the duration of exposure resulting from tidal height, increased temperatures resulting from seasonality, and marsh plant density. Burrows were deeper and more expansive at both higher tidal levels and higher temperatures in summer. Burrow dimensions were sharply reduced with the disappearance of adult macrofauna in areas where the roots of the marsh plant Suaeda japonica were dense. The significance of this study lies in quantifying the burrow dimensions of the entire macrofaunal community, rather than just a single population, and confirming their spatial and temporal variation with respect to physical conditions of the habitat. Environmental factors responsible for variation in burrow dimensions are discussed.