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.     

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.     

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.     

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.     

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.     

The Biology of Upogebia pusilla (Petagna) (Decapoda, Thalassinidea) I. The Burrows

Abstract. The biology of the thalassinidean shrimp Upogebia pusilla was studied on a tidal flat in the Lagoon of Grado (Northern Adriatic). Burrows were investigated using in situ resin casting and with additional in situ and laboratory observations. Burrows show a basic pattern consisting of a U or a double U with turning chambers and a vertical shaft. Mean burrow diameter depends on animal size, it is smaller than the rigid carapace of the animal. Dimensions, distance between openings, depth of U, total depth, volume and surface are size dependent. The burrow wall is smooth and oxidized. Burrows are always inhabited by a single shrimp. Although they overlap, they are never interconnected. They are mainly constructed by compression of the sediment and are relatively permanent structures. A comparison of the burrows of Thalassinidea with regard to shape, number and appearance of openings, dimensions, properties of the burrow wall and dynamics is given.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

青岛潮间带底内蟹类生态及洞穴观察

栖息于潮间带的海生动物中,营底内生活的种类占很大的比例。这类动物以独特的生活方式适应或克服潮间带的各种不利条件而得到生存和繁衍。从报道来看现代海洋底内动物泂穴的研究具有多方面的意义。如某些潮间带底内动物的鉴定,仅以外部形态作依据显然不够,还需以其他生态资料,如粪便、洞穴、卵块等作参考。有些潮间带动物的地理分布范围与其营底内生活有关，它们可以借助洞穴的保护而渡过寒暖不适的气候。更重要的是这方面研究能对古生态学和地史学问題的探讨提供科学依据。有很多种潮间带底内动物洞穴、排出物和爬行痕迹等被作为“生活遗迹”而保存在化石中,因此这类化石的沉积岩层实际上已成为该类动物生存发展年代的记载。但是,在岩层古生态学分析研究中,比较困难的是如何区别古代生物居住、死亡或埋藏的地点,这个同题涉及到对沉积物形成的地点和过程的确定,进一步就可用这些资料来研究古代海岸变迁和海面的升降。就潮间带底内动物的居住和死亡地点(埋藏与死亡地点也可以不同)来说,有的可能一致(如某些不太活动的种类),有的则不一致(如贝类)，作为古生态学依据,各具其长短,贝类无洞穴化石,其贝壳可成为化石,但埋藏与生活地点往往不一致;蟹类的优点是洞穴能成化石，其上并留有爬行痕迹;洞穴化石的埋藏与动物原来生活地点一致(亦有个別情况不一致)。早在地史年代就已有不少虾蟹类遗体及生活迹迹化石被发现:因此，现代潮间带蟹类洞穴的研究不仅对认识或鉴定古代甲売类动物有价值,而且对进一步鉴定种类、寻找演化线索和促进古生态学的发展都具有重要意义。 青岛潮间带在我国北方沿岸潮间带中颇具代表性。潮汐是规则的半日潮。 著者曾于1954-1964年,选以下四个点进行了调查:1,沧口潮间带,位于胶州湾内东北区中部,座东向西，坡度小的泥质和泥沙质滩涂;2.沙子口潮间带,位于青岛以东,为沙质滩涂;3.湛山湾潮间带:位青岛以东近郊,为沙质滩涂;4.大黑澜潮间带,位于胶州湾口,为岩石岸滩。每月进行二次大潮采集,每一采样点取两个定量(50×50厘米)样品和若于定性补充样品。从1963年5月至1964年9月,结合甲壳类生态进行了专题调查,本文就是根据上述调查,在软底质滩涂所获11种蟹类材料总结的,对所见各种蟹类的洞穴均经现场观察、测量并作了详细记录和拍照。对同一种蟹类洞穴的形态在不同地点和时间内所出现的变化也给予了特别的注意。 本文定稿过程中征得了山东海洋学院李嘉泳及本所曾呈奎,刘瑞玉、郑执中、吴宝铃、齐钟彦、马绣同等同志的修改意见,文中插图、照片由王兴虞、宋华中同志给予帮助,在此一并表示感谢。     

Burrow morphologies in north-central Pacific sediments

An examination of split core photographs from the north-central Pacific reveals the presence of three different burrow morphologies. Simple burrows occur throughout the region. Halo burrows are present in water deeper than 4,000 m (3,000 m along the equator) and removed from continental areas. Imbricate burrows are uncommon and have an irregular distribution. Organisms responsible for the burrows are interpreted as surface to near-surface feeders which do not penetrate deeply into the sediments. Thus, the zone of sediment mixing caused by burrowing ranges from 10 to 30 cm and averages 20 cm.     

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.     

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.     

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.