Impact of fiddler crab foraging and tidal inundation on an intertidal sandflat: season-dependent effects in one tidal cycle

Intertidal sandflats inhabited by fiddler crabs are ideal systems in which to study the effects of physical and biological processes. This study addressed two questions: (1) Do fiddler crab feeding and tidal inundation have measurable effects on the sandflat over one tidal cycle? (2) Does the sandflat change over the course of a year? In field exclusion/inclusion experiments, fiddler crabs reduced sediment organic content by 40%, Chlorophyll a levels by 20% and meiofaunal density by 60% in one tidal cycle. Effects were most pronounced in the spring when organic content and meiofaunal densities were maximal. Effects of foraging were not erased by the tide and accumulated over time. The sandflat had highest levels of all variables in spring and minimal levels during summer and fall. Crabs graze the sandflat to minimum levels in the spring. Due to crab foraging, the flat is barren during the summer and fall, and recovers during the winter when crabs are minimally active.     

Habitat partitioning between prey soldier crab Mictyris brevidactylus and predator fiddler crab Uca perplexa

Interaction and habitat partition between the soldier crab Mictyris brevidactylus (prey) and the fiddler crab Uca perplexa (predator) were examined at a sandy tidal flat on Okinawa Island, Japan, where they co-occur. Both live in dense colonies. When the soldier crabs were released in the densely populated habitat of the fiddler crab, male fiddler crabs, which maintain permanent burrows in hard sediment, preyed on small soldier crabs and repelled large ones. Thus, the fiddler crabs prevented the soldier crabs from trespassing. It was also observed whether soldier crabs burrowed successfully when they were released 1) where soldier crab burrows just under the sand were abundant, 2) in a transition area between the two species, 3) an area without either species, and 4) where artificial tunnels simulated soldier crabs' feeding tunnels were made by piling up sand in the area lacking either species. In contrast to the non-habitat area, many soldier crabs burrowed in the sediment near the release point in the tunnel, transition and artificial tunnel areas. This indicates that the feeding tunnels on the surface attracted other crabs after emergence. When the large male fiddler crabs were transplanted into the artificial burrows made in soft sediment of the soldier crab habitat, all left their artificial burrows by 2 days. In the fiddler crab habitat, however, about one-third of the transplanted male fiddler crabs remained in the artificial burrows after 3 days. The soldier crabs regularly disturb the sediment by the up and down movement of their burrow (small air chamber) between tides. This disturbance probably prevents the fiddler crab from making and occupying permanent burrows. Thus, it appears that these crabs divide the sandy intertidal zone by sediment hardness and exclude each other by different means.     

Coupling between microphytobenthic biomass and fiddler crab feeding

Among the organic matter ingested by fiddler crabs, microphytobenthos is of fundamental importance because it is their main N source. Microphytobenthos abundance generally develop semilunar changes as the dynamics of tidal exposures and day-night cycle are not held constant across days, modifying the balance between growth and mortality. In this study we explored the coupling between temporal dynamics in microphytobenthos abundance and crab feeding activity. We measured the Chlorophyll a content in the 2 mm surficial sediment surrounding the burrows and the crab feeding activity over two semilunar cycles. Chlorophyll a and crab feeding activity showed biweekly cyclic dynamics. Crabs did not concentrate feeding activity around days with maximum abundance of microhytobenthos. This phase difference between both dynamics could be the result of the crab feeding impact, but a crab experimental exclusion showed that the temporal dynamics of Chlorophyll a content stayed unchanged when feeding activity was removed. Comparisons between fed and unfed sediment suggest that the feeding efficiency changes with tidal dynamic. Crabs achieved more than 50% of Chlorophyll a extraction during days of highest feeding activity, and less than 30% during days of low feeding activity or low microhytobenthos abundance. Furthermore, comparisons of fed sediment between consecutive days indicated that Chlorophyll a was completely replenished during days with high flooding tides, but partially replenished during days near neap tides. Environmental conditions affecting feeding efficiency may select crabs to concentrate feeding activity before days with the highest microhytobenthos abundance. The low feeding impact on microphytobenthos dynamics suggests that fiddler crabs would not control microhytobenthos abundance and thus unable to absorb the increasing eutrophication of studied estuarine areas.     

Experimental studies on the foraging behavior of the sand fiddler crab Uca pugilator (Bosc, 1802)

The purpose of this study was to test the hypothesis that foraging sand fiddler crabs. Uca pugilator (Bosc), move through the habitat in response to low substratum food levels even though these movements may take the crabs considerable distances from the safety of the burrow area. Chl a and ATP concentrations were used as measures of food density in foraged and unforaged substratum. Field and laboratory feeding experiments showed that crab foraging intensity in a habitat patch was directly correlated with food density in the patch either in the presence or absence of alternative food patches. Other experiments showed that sand fiddlers can respond to differences in food level on a scale of millimeters and do this by probing the substratum with minor chelae. Food levels in aggregations of non-ingested particles harvested by sand fiddlers, feeding pellets, correspond to low foraging intensities predicted from foraging experiments and crabs exhibit low foraging intensities on substratum patches derived from feeding pellets. Substratum food levels in two distinct areas corresponded to high predicted foraging intensities and there was no consistent trend in the level of food in the burrow vs. the nonburrow microhabitats. These results suggest that the movements of foraging sand fiddlers are to some extent controlled by the reduction in substratum food levels due to feeding during a single foraging episode. Sand fiddlers can extract over 70% of the food from harvested substratum over a broad range of substratum food densities but harvest only 42% of the available substratum.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

Comparative Burrow Architectures of Resident Fiddler Crabs (Ocypodidae) in Indian Sundarban Mangroves to Assess Their Suitability as Bioturbating Agents

Excavation of burrows by fiddler crabs (genus Uca) is an important component in mangrove ecosystem functioning. This bioturbation activity can be measured by analysing the burrow architecture of these crabs. The aim of the present study is to describe and evaluate inter specific differences in the burrow morphologies of four species of fiddler crabs (Uca rosea, Uca triangularis, Uca dussumieri and Uca vocans) using polyester resin casts of the burrows. For each of the species, sex and carapace width (CW; mm) were determined for all the individuals. Three burrow morphological characters viz. burrow diameter (BD; mm), total burrow depth (TBD; mm) and burrow volume (BV; cm³) were considered during the study. Density of each species throughout the year was also assessed. For all the species BD and BV were higher in case of males compared to the females and they showed significant positive correlation with the CW of the burrow inhabitants. The amount of sediment excavated by each crab was evaluated in terms of BV. Among all the studied species, U. rosea was established as the most potent bioturbative candidate in the studied mangrove due to their greater density and moderate ability to excavate burrow.     

The Contribution of Crab Burrow Excavation to Carbon Availability in Surficial Salt-marsh Sediments

Geomorphology, vegetation and tidal fluxes are usually identified as the factors introducing variation in the flushing of particulate organic matter (POM) from tidal marshes to adjacent waters. Such variables may, however, be insufficient to explain export characteristics in marshes inhabited by ecosystem engineers that can alter the quantity and quality of POM on the marsh surface that is subject to tidal flushing. In this study we evaluated the balance between transfer of buried sedimentary organic carbon (C) to the marsh surface due to crab excavation (measured from the mounds of sediment excavated from burrows) and outputs of C from the surface due to sediment deposition within crab burrows (estimated from sediment deposited within PVC burrow mimics), in a Southwestern Atlantic salt marsh supporting dense (approximately 70 ind m⁻²) populations of the crab Chasmagnathus granulatus. C excavation by crabs was much greater than deposition of C within crab burrow mimics. Per area unit estimates of the balance between these two processes indicated that crabs excavated 5.98 g m⁻² d⁻¹ and 4.80 mg m⁻² d⁻¹ of total and readily (10 d) labile C, respectively. However, sediments excavated by crabs showed a significantly lower content of both total and readily-labile C than sediment collected in burrow mimics. This indicates that ecosystem engineering by burrowing crabs causes a net decrease in the concentration of C in the superficial sediment layers and, thus, an overall decrease in the amount of C that can be washed out of the marsh by tidal action. Incorporating the in situ activities of ecosystem engineers in models of marsh export should enhance understanding of the function of marshes in estuarine ecosystems.     

Composition and abundance of shorebird and macroinvertebrate communities differ according to densities of burrowing fiddler crabs in tropical intertidal flats

The burrowing activity of fiddler crabs inhabiting intertidal flats creates visually distinct patches within these habitats. However, differences in the composition and abundance of shorebirds and their macroinvertebrate prey between areas inhabited or not by crabs are yet to be studied. Here, we compare the macroinvertebrate and shorebird assemblages in low and high crab density areas in the intertidal flats of the Bijagos archipelago, Guinea-Bissau. High crab density areas are associated with lower richness and densities of macroinvertebrates. Shorebird assemblages were also less rich at high crab density areas and the differences in species composition occurred according to prey type preferences. Fiddler crab density was the most important variable explaining macroinvertebrate abundance, after accounting for the effects of fine fraction of sediment and distance to coast. Nonetheless, a controlled experimental setup would be required to attribute differences found to the engineering activity of fiddler crabs rather than other unaccounted habitat features. Our findings suggest that crab patches should be taken into account when assessing the distribution and abundance of macroinvertebrates and shorebirds in intertidal areas. Since low and high crab density areas differ markedly in terms of shorebird carrying capacity, monitoring variations in their extent will be important to interpret past and present population trends.

     

Burrow structure and use in the sand fiddler crab, Uca pugilator (Bosc)

Uca pugilator, the sand fiddler crab, constructs two kinds of burrows in protected, sandy upper-intertidal and supratidal substrates on the west coast of Florida. Temporary burrows are built and used as a refuge by non-breeding crabs during high tide periods and at night when crabs cease feeding in the intertidal zone. Breeding burrows are constructed and defended by courting males and are the site of mating, oviposition and the incubation of eggs by females. Up to three ovigerous females may be accommodated in a single breeding burrow, each female sequestered in a separate terminal chamber. The construction and defence of burrows specialized for breeding may be an adaptive response by males to the preferences females exhibit when selecting a breeding site.     

Does the presence of the SW Atlantic burrowing crab Chasmagnathus granulatus Dana affect predator–prey interactions between shorebirds and polychaetes?

The burrowing crab Chasmagnathus granulatus is an important bioturbator that generates dense burrow assemblages (crab beds) characteristic of intertidal habitats of SW Atlantic estuaries. Crab bioturbation affects the topography and hydrodynamics of the sediment, increasing sediment water and organic matter content, decreasing sediment hardness and changing the grain size frequency distribution. In this study, we found that burrowing crabs can decrease the impact of predation by shorebirds on polychaetes. The polychaete Laeonereis acuta Treadwell has U-shaped burrows outside crab beds, which are associated with surface deposit-feeding while their burrows are mainly I-shaped inside which is associated with subsurface deposit feeding behavior. This pattern is likely the result of larger vertical sediment mixing inside crab beds due to crab burrowing. As a result of their feeding strategy, polychaetes appear on the surface more often outside crab beds, which increases their availability for shorebirds. In addition, shorebird species differentially use crab beds. The White-rumped Sandpiper, Calidris fuscicollis Vieillot, preferentially forage outside crab beds, meanwhile the Two-banded Plover Charadrius falklandicus Latham forage more frequently inside crab beds. However, experiments excluding shorebirds inside and outside crab beds showed negative effects of shorebirds only outside crab beds. Thus, our results show that the SW Atlantic burrowing crab C. granulatus affects the strength of the predator–prey interaction between shorebirds and polychaetes.     

Impact of fiddler crabs (Uca spp.) on rates and pathways of benthic mineralization in deposited mangrove shrimp pond waste

The impact of fiddler crabs (Uca spp.) on benthic mineralization rates and pathways in deposits of shrimp pond waste (SPW) with planted mangrove trees (Rhizophora apiculata) were determined in the Ranong mangrove forest, Thailand. Sediment metabolism, measured as CO₂ flux, increased by 2- to 3-fold when either fiddler crabs or mangrove trees were present compared to control plots. Sulfate reduction rates (SRR) were always high and partitioning of various electron acceptors to total carbon oxidation revealed that sulfate reduction contributed by >90%, with iron reduction being important only near the sediment-water interface. However, significant iron reduction appeared down to 7 cm when bioturbation and plant roots were combined in the easily oxidizable mangrove sediment (MS), indicating that infaunal activity and plant roots were able to alter the substratum. Microprofiles around individual Uca burrows showed 46% lower SRRs in a 15-mm-thick oxidized layer around the burrows compared to the surrounding sediment. The burrow wall environment appears to be a zone of intense reoxidation of reduced compounds as indicated by low pools of reduced sulfide compounds and a high Fe(III) content. Despite the decreased SRRs near the burrow, and the introduction of Fe(III) to deeper sediment layers, fiddler crabs and mangrove trees have only limited impact on rates and partitioning of anaerobic carbon mineralization in the SPW. This lack of response was attributed to the relative small volume of sediment affected by crab activities.     

Invertebrate perception: measuring depth intervals through path integration and vision

A foraging fiddler crab can estimate how close a potential intruder is from its burrow entrance, even when the entrance in the sand is invisible to the crab. Recent work shows that, to assess this depth interval, crabs combine information from vision and path integration in an unusual manner.     

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance (''surface mating'') or leave their own burrow and sequentially enter and leave (''sample'') courting males'' burrows, before staying in one to mate underground (''burrow mating''). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.     

Role of fiddler crabs of a subtropical intertidal flat on the fate of sedimentary fatty acids

Fatty acid compositions of three ocypodid crabs, Uca vocans, U. lactea and Scopimera globosa, collected on a subtropical intertidal flat (Okinawa, Japan), were analysed as well as the surrounding surface sediment of their burrows, their food pellets and their faeces. U. vocans and U. lactea exhibited a higher concentration of fatty acids than S. globosa. Food pellets revealed a higher relative contribution of saturated fatty acids and lower amounts of polyunsaturated and branched fatty acids than in the surface sediment samples. These differences of fatty acid compositions suggest an initial selection of food before the particle sorting in the buccal region. This result, and also because of the relatively high fatty acid content of faeces, indicate that fiddler crabs play a significant role in the fate of these lipids in the subtropical intertidal flats. Comparative analysis of the fatty acid compositions of the different samples indicates that these fiddler crabs actively sorted bacteria in order to ingest them. Diatoms seem to be used in larger amount by U. vocans may be because of its higher mobility while looking for food. Other primary producers present in the environment, mangrove leaves and algae, have been sorted but contributed weakly to the organic matter pool of the surface sediment and therefore to the diet of the crabs.     

The validity of burrow counting in estimating the densities of Chiromantes dehaani and Ilyoplax deschampsi in tidal flat at Yangtze Estuary

It has been confirmed that the crabs play significant roles in the structure and function of coastal wetland ecosystems, such as mangroves and salt marshes. However it is not easy to estimate the abundance and density of burrowing crabs effectively, thus a further understanding of roles of crabs in these ecosystems has been lagged. Some studies have discussed the suitability of several census techniques, such as burrow counting method in estimating crab density in mangroves. The validities of burrow counting method and other census techniques in estimating crab density, however, has not been tested in salt marshes, especially where vegetation are dense. In this study, we tested the validity of burrow counting method in estimating the densities of Chiromantes dehaani and Ilyoplax deschampsi in tidal flat with dense vegetation of Phragmites australis and Zizania aquatica at Yangtze Estuary through comparing densities estimated by the burrow counting method and the excavation. Burrow counting averagely underestimated the density of C. dehaani by 15% and the degree of underestimation varied among vegetations and habitats (from an overestimate by 23% to underestimate by 41%). Burrow counting averagely overestimated the density of I. deschampsi by 43% and the degree of overestimate varied from 0% to 133% depending on the vegetations and habitats. The percentage of occupied burrows and the number of crabs sharing one burrow were important factors influencing the validity of estimating crab density through burrow counting method.     

The validity of burrow counting in estimating the densities of Chiromantes dehaani and Ilyoplax deschampsi in tidal flat at Yangtze Estuary

It has been confirmed that the crabs play significant roles in the structure and function of coastal wetland ecosystems, such as mangroves and salt marshes. However it is not easy to estimate the abundance and density of burrowing crabs effectively, thus a further understanding of roles of crabs in these ecosystems has been lagged. Some studies have discussed the suitability of several census techniques, such as burrow counting method in estimating crab density in mangroves. The validities of burrow counting method and other census techniques in estimating crab density, however, has not been tested in salt marshes, especially where vegetation are dense. In this study, we tested the validity of burrow counting method in estimating the densities of Chiromantes dehaani and Ilyoplax deschampsi in tidal flat with dense vegetation of Phragmites australis and Zizania aquatica at Yangtze Estuary through comparing densities estimated by the burrow counting method and the excavation. Burrow counting averagely underestimated the density of C. dehaani by 15% and the degree of underestimation varied among vegetations and habitats (from an overestimate by 23% to underestimate by 41%). Burrow counting averagely overestimated the density of I. deschampsi by 43% and the degree of overestimate varied from 0% to 133% depending on the vegetations and habitats. The percentage of occupied burrows and the number of crabs sharing one burrow were important factors influencing the validity of estimating crab density through burrow counting method.     

Morphological and behavioural adaptations to the rocky substrate by the fiddler crab Uca panamensis (Stimpson, 1859): preference for feeding substratum and feeding mechanism

The fiddler crab Uca panamensis (Stimpson, 1959) inhabits rocky shores. We examined its preference for feeding substratum—sand or rock—and its manner of feeding. The crab made its burrow in the sand among rocks but preferred to feed on rocks. The feeding time decreased as the distance between the burrow and the rock increased. We consider this to be a result of exclusive interaction among the crabs because they defended their feeding area on the rocks against others.The crab wetted a small area of rock with water held in the branchial chambers before and during feeding. It pinched up the wetted surface in the minor chelipeds, which have bundles of setae on the posterior tips of the dactyl and pollex, and put the material into its buccal cavity. It never expelled sand pellets while feeding on rock, which indicates that it swallowed the food particles directly, without sorting. The bundles of setae retained water by capillary attraction, which suggests that they capture the suspended fine food particles scraped from the rock. The wetting action may prevent the fine materials from dispersing. We consider that morphological alteration of the minor chelipeds, the application of water from the branchial chambers, and direct swallowing permit the fiddler crab to feed on fine materials attached to rocks.     

Why do fiddler crabs build chimneys?

Chimneys are mud mounds built by fiddler crabs that encircle the entrance to their burrow. Their function in many species is unknown. In Uca capricornis, crabs of both sexes and all sizes build chimneys, but females do so disproportionately more often. There are no differences in the immediate physical or social environments between crabs with and without a chimney. Chimney owners spend less time feeding and more time underground than non-owners. We show experimentally that burrows with a chimney are less likely to be located by an intruder. It is possible that some crabs construct chimneys around their burrow to conceal the entrance and reduce the risk of losing it to an intruder.     

长江口芦苇和互花米草盐沼湿地蟹类洞穴分布特征及主要影响因子

蟹类洞穴是蟹类在潮间带盐沼生存、繁衍的特征性结构,具有重要的生态功能。洞穴分布特征及其影响因子的分析,是深入探讨蟹类及其洞穴的生态系统功能的重要基础。2015年10月,在崇明北滩单一芦苇(Phragmites australis)群落,单一互花米草(Spartina alterniflora)群落和芦苇-互花米草混合群落3种典型生境中,对蟹类洞穴的分布特征及其相关的大型底栖动物、植被、沉积物等的特征参数进行了调研与分析。结果表明,生境类型差异对蟹类洞穴分布特征及相关生境因子具有重要影响。蟹类洞穴的分布密度和开口直径在不同生境间存在显著差异(P0.05),且单一芦苇群落生境内洞穴密度要显著高于单一互花米草群落生境(P0.05),洞穴开口直径在单一互花米草生境要显著高于单一芦苇生境(P0.05);大型底栖动物生物量、密度、植物地下部分生物量在不同生境间差异不显著(P0.05),而植株密度、活植株高度、植物地上部分生物量以及沉积物含水率、p H、氧化还原电位在不同生境间存在显著差异(P0.05)。沉积物中值粒径,总氮含量和总碳含量在不同生境间的差异随深度不同会发生变化。不同生境主要生境因子的差异是导致蟹类洞穴分布特征不同的根本原因;蟹类洞穴分布特征受多个生境因子的综合作用。筛选的生境因子的组合虽然与洞穴分布特征具有显著相关性,但相关系数较小。未来研究中需要拓展生境因子涵盖范围,加强多因子综合作用分析。