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.     

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.     

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.     

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 Biology of Upogebia pusilla (PETAGNA) (Decapoda, Thalassinidea)

Abstract. The distribution of the thalassinidean shrimp Upogebia pusilla was studied at four sites in the North Adriatic Sea: 1) a tidal flat in the lagoon of Grado, 2) a tidal flat at Lido di Staranzano near the mouth of the Isonzo, 3) a mud flat seawards of a salt marsh in a protected bay north of Rovinj, and 4) a sublittoral station in 6m depth near Aurisina. Information on grain size distribution, organic content, amount of debris, redox profiles and pH of sediment as well as temperature and salinity is given. Density, as determined by hole counts and a hole: burrow relationship derived from resin casting, generally increased with increasing water depth in the intertidal. The upper limits ranged between +10cm (Rovinj) and -20cm (Grado); densities between mean water and low water level varied strongly due to microtopography and macrophyte cover. Maximum densities in the intertidal ranged from 189 (Grado) to 2420 (Rovinj, juveniles) animals m². Shrimp density at the sublittoral station ranged between 80 and 230 m"². The zonation of the Upogebiidae and Callianassidae with respect to environmental parameters is discussed.     

Population biology of the fiddler crab Uca annulipes (Brachyura: Ocypodidae) in a tropical East African mangrove (Mozambique)

The community structure of the tropical fiddler crab Uca annulipes (H. Milne Edwards) has been studied in Mozambique for the first time. Ten 0.25 m² squares were randomly sampled on a monthly basis during low tide periods from January to December 2002 at Costa do Sol Mangrove, Maputo Bay, southern Mozambique. A total of 981 crabs were captured, of which 494 were males (50.36%), 231 non-ovigerous females (23.85%) and 253 ovigerous females (25.79%). The overall size frequency distribution was unimodal, with males being larger than both ovigerous and non-ovigerous females. Females were more abundant in the smallest size classes while males outnumbered females in the largest ones. The overall sex ratio (1:0.99) did not differ from the expected 1:1 proportion, but significant deviations were observed in January, April, August and November. Results indicate that the size frequency distributions of this species can also be determined through direct analysis of burrow openings. Breeding took place year-round with two peaks of spawning in summer (January and December). Recruits were present throughout the year, with high abundance in summer probably due to the high reproductive activity observed in this season. The present results may allow us to suggest that U. annulipes follows a rapid breeding cycle accompanied by a rapid larval development and settlement in the study area.     

The Biology of Upogebia pusilla (PETAGNA) (Decapoda, Thalassinidea)

Abstract. Populations of the thalassinidean shrimp Upogebia pusilla were studied on tidal flats in the Northern Adriatic Sea. Biometric analysis showed a sexual dimorphism, especially in propodus size. Size frequency distributions revealed the presence of large animals during all seasons; recruitment by juveniles occurred in autumn. Moult intervals and moult increments were determined in the laboratory and used to generate growth curves which were compared with those calculated from size frequency distributions. The life span of U, pusilla is over 5 years. Ovigerous females occurred between March and September. Egg numbers were high in spring, lower in summer and increased with body size. The incubation time of embryos was 35 days; a female produces an estimated number of three egg batches during the breeding season. A total annual production of 994 kJ was estimated for a theoretical population of 100 animals; 13.5 % is spent for somatic growth, 31 % for egg production of females, and 55.2% is lost as exuviae. The population structures, growth and breeding patterns, as well as embryonic and larval development within the Upogebiidae and Callianassidae are discussed.     

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.     

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.     

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.     

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.     

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.     

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.     

The role of crabs (Macrophthalmus japonicus) burrows on organic carbon cycle in estuarine tidal flat,Japan

The objective of this study is to elucidate the burrow structure and to clarify the role of burrows in material cycle in the tidal flat. In our work, we focused on the dominant species in muddy tidal flat, crab Macrophthalmus japonicus.Burrow structure of Macrophthalmus japonicus was investigated on a Katsuura river tidal flat in Tokushima prefecture, Japan, using in situ resin casting. Sampling was conducted in August 2006, and a total of 48 burrow casts were obtained. Burrows consisted mainly of J-shaped structures (98%) while the rest belonged to U-shaped structures (2%). The maximum measured burrow volume was 120 cm³ and wall surface area was 224 cm², while maximum burrow length and depth were 23.2 cm and 16.5 cm, respectively. Burrow volume and surface area were strongly correlated with carapace width of M. japonicus. Investigation of the individual number of M. japonicus in 13 quadrats (50 × 50 × 20 cm) was conducted using 2 mm sieve. The number of M. japonicus was 15–31 ind./m². Using cohort analysis we estimated that surface area of burrows was 0.07–0.15 m²/m².CO₂ emission rate was measured at the surface sediment during the period from June to December 2008. Results varied from 13.8 ± 2.2 to 49.4 ± 3.2 mg CO₂/m²/h, and organic carbon decomposition was 3.8 ± 0.6–13.5 ± 0.9 mg C/m²/h. This leads the increase of organic carbon decomposition by 1.1 times, because of the expansion of the tidal flat surface area by burrowing activity. Organic carbon decomposition in burrow walls therefore contributed to organic matter decomposition in the tidal flat. These results indicated that in situ activities of Macrophthalmus japonicus significantly influence the material cycle and it is important to consider the existence of burrow in order to understand the fluxes of materials and to evaluate the purification function of the tidal flat.     

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.     

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.     

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.     

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.     

Catch, survey and life-history data for shrimp (Pandalus borealis) off Jan Mayen

The Jan Mayen area has an extreme environment with low temperatures and infrequent, but abrupt temperature changes. The shrimp population here is considered to be on its edge of distribution. The life-history parameters are in the same range as in other high-latitude shrimp populations and are characterized by slow growth, large size at maturation and extended longevity. Irregular and sporadic commercial exploitation limit fishing mortality and give the population life-history parameters not previously seen in other areas. The Jan Mayen shrimp are large compared to, e.g., the Barents Sea shrimp and can reach a maximum carapace length (L_max) of 37 mm and an age of 10–11 years. The large size at sex transformation (L₅₀, >24 mm) and analyses of length–frequency distributions indicate that the shrimp may be 6–7 years of age before changing sex. The change in L_max and L₅₀ observed during the study period is probably caused by increased natural mortality due to sudden temperature changes or due to increased predation, rather than increased growth rates. The life-history strategy of shrimp in the Jan Mayen area can be explained by factors such as depth, temperature and population density variations caused by fluctuation in recruitment and mortality.The shrimp fisheries in the Jan Mayen area began in the late 1970s and reached an annual landing of 2000 tonnes in 1985, and since then landings have oscillated around 500 tonnes depending on a combination of factors. The survey indices of stock biomass varied between 3000 and 6600 tonnes. For most years, the highest shrimp densities are at a depth of 200–299 m, while large shrimp (and therefore also female shrimp) are dominant at depths greater than 300 m.Fish community data were studied as the composition of the demersal fish community is an integrated response to environmental conditions and as predation affects the shrimp stock. Polar cod and capelin are the most abundant fish species in the study area. A high number of blue whiting was registered in 1979, but the number declined in 1980 and 1981 as temperature decreased. During the surveys in 1994 and 1995, no blue whiting was registered. A few individuals were found again in the 1999 samples. The number of Greenland halibut has declined from the beginning of the 1980s to the 1990s.     

Rejection of Intracoelomic Invading Material by Leptosynapta inhaerens (Echinodermata: Holothuroida): A Process of Ecological Significance?

Abstract. The synaptid holothuroid Leptosynapta inhaerens has the ability to expel unwanted particles that enter its body cavity. Intracoelomic particles ( viz. experimentally injected carmine particles) are trapped either by a coelom-produced mucoid net or by specialized organs (the vibratile urnae) that occur in bands in some interradial areas. Whatever the trapping method, particles are incorporated into dense mucoid masses that move towards the posterior part of the body cavity, in the vicinity of the rectum. These aggregates then cross the rectal wall through rectal pores, mix with the faeces, and are eventually incorporated into the wall of the synaptid burrow. Clearance of foreign particles requires 2 to 3 days. The coelom-cleaning system of synaptids appears to be the most effective within the class Holothuroidea , being well designed for endofaunal organisms whose walls are delicate and easily rupture.