Adult fiddler crabs Uca pugnax (Smith) enhance sediment-associated cues for molting of conspecific megalopae

Adult-associated chemical cues can stimulate settlement and metamorphosis of invertebrate larvae into habitats with an enhanced likelihood of juvenile and adult survival. For example, sediments from adult fiddler crab habitat stimulate fiddler crab megalopae to metamorphose (molt) sooner than sediments without adult cues. A similar stimulation of molting occurs after exposure to waterborne chemical cues from adult habitats and to exudates and extracts of adult crabs. We tested whether sediments from habitats without adult Uca pugnax (Smith), which do not stimulate molting of their megalopae, could become stimulatory through brief exposure to adult crabs. Sediments were collected from tidal flats at several distances (∼ 1 m, ∼ 50 m, and ∼ 5.4 km) from adult habitats, and incubated for 24 h with or without adult crabs. Molting rates of laboratory-reared megalopae exposed for 48 h to adult-conditioned sediments were compared to those for untreated controls. Sediments collected in or within 1 m of adult habitat elicited the highest molting rates, and natural sediments from 50 m and 5.4 km had little or no effect on molting. However, incubating sediments collected away from adult habitat with adult crabs produced a higher molting response, and the magnitude of the enhancement increased with distance from adult habitat. Results suggest that the chemical cues that adult crabs release are retained by sediments and consequently stimulate molting of megalopae, regardless of the nature of the sediments themselves. Lack of chemical cues may retard colonization of newly created or heavily disturbed habitats that are otherwise suitable settlement and adult habitat.     

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.     

The SW Atlantic burrowing crab Chasmagnathus granulatus Dana affects the distribution and survival of the fiddler crab Uca uruguayensis Nobili

In this paper, we address the question of whether the presence of the burrowing crab Chasmagnathus granulatus affects the habitat use of the fiddler crab Uca uruguayensis. Field samples showed that the species have a disjoint spatial distribution. Male fiddler crab density decreased in zones with C. granulatus, however, female density increased. Male fiddler crabs avoided feeding on sediment affected by C. granulatus and were more preyed. Predation was higher during the fiddler crab reproductive season and, probably due to predation risk, males showed lower reproductive display in shared zones. Field experiments shows that when C. granulatus were excluded, densities of U. uruguayensis increased mainly due to an increase in density of males. Habitat differentiation of these species may be because C. granulatus affects U. uruguayensis in several ways, including direct predation, disturbance and behavioural changes associated to predation risk. Males and females are affected differentially probably because of the extreme sexual dimorphism of this crab species. Coloration on enlarged claw and waving activities are all factors that increase predation risk for male and the presence of only one feeding claw may increase sediment-mediated effects.     

Entrainment of the activity rhythm of the mole crab Emerita talpoida

The mole crab Emerita talpoida migrates with the tide in the swash zone of sand beaches. A circatidal rhythm in vertical swimming underlies movement, in which mature male crabs show peak swimming activity 1-2 h after the time of high tides at the collection site. In addition, there is a secondary rhythm in activity amplitude, in which crabs are maximally active following low amplitude high tides and minimally active following high amplitude high tides. The present study determined the phase response relationship for entrainment of the circatidal rhythm with mechanical agitation and whether the cycle in activity related to tidal amplitude could be entrained by a cycle in the duration of mechanical agitation at the times of consecutive high tides. After entrainment with mechanical agitation on an orbital shaker, activity of individual crabs was monitored in constant conditions with a video system and quantified as the number of ascents from the sand each 0.5 h. Mechanical agitation at the times of high tide, mid-ebb and low tide reset the timing of the circatidal rhythm according to the timing relationship to high tide. However, mechanical agitation during flood tide had no entrainment effect. In addition, a cycle in duration of mechanical agitation entrained the rhythm in activity amplitude associated with tidal amplitude. Both rhythms and entrainment effectiveness over the tidal cycle may function to reduce the likelihood of stranding above the swash zone.     

Predator-driven fragmentation of fiddler crab droves into selfish miniherds of biased composition

An explanation for animal groups is the selfish herd, characterized by aggregation as each member tries to shield itself from a predator by moving into a tight gap between other members. We test the hypotheses that: (1) droves, the large feeding groups of fiddler crabs, are selfish herds; (2) the miniherds that form when droves fragment on approach of a large predator are selfish herds; (3) selfish herds form when refugia are unlikely to be reached before an approaching predator arrives; and (4) the composition of selfish miniherds is biased toward individuals most vulnerable to predation. The study was conducted in South Carolina (USA) by videotaping the movements of sand fiddler crabs Uca pugilator when approached by a human predator. In both droves and miniherds, interindividual distance decreases with predator approach, consistent with behavior in a selfish herd. However, two other expectations for selfish herds—herd cohesion and sacrificing distance from the predator in order to get closer to other herders—are only met in miniherds. Crabs farther from refugia are more likely to form and remain in miniherds, indicating that selfish herding is only favored when refugia cannot be quickly reached. The composition of the smallest miniherds, consisting of 2-18 crabs, is biased toward females and small males. These individuals may be more vulnerable to predation because they lack the enlarged claw of large males that deters some predators. The small miniherds are relatively homogeneous with respect to the size and sex of their members, which may enhance cohesion and effectiveness as selfish herds. Miniherds will be effective selfish groups when predator attack has a significant vertical component and when the strike distance is large relative to both the size of the prey and the distance between group members. Droves are not selfish herds but permit crabs to flee feeding grounds as members of selfish miniherds.     

The relative importance of substratum characteristics and recruitment in determining the spatial distribution of the fiddler crab Uca uruguayensis Nobili

The roles of sediment characteristics and the pattern of recruitment in influencing the abundance of the fiddler crab Uca uruguayensis on Argentinean mudflats were evaluated. The density of adult crabs showed a patchy distribution related to the sediment thickness (depth at which a layer of fossil shells are buried), but the density of juvenile crabs was not coupled with the density of adult crabs. In a field experiment, fossil shells were removed and the density of crabs significantly increased, which demonstrates that the presence of the layer of shells is a structure that may hinder the establishment of burrows. The density of crabs was related to sediment thickness, sediment torque and organic matter content. The importance of each of these variables was different for adult and juvenile crabs, indicating that the distribution of adult crabs may be caused by mechanisms affecting adult crabs themselves and is not established by the recruitment pattern. Moreover, in a field experiment, the density of juveniles decreased when adult crabs were added, and increased when adult crabs were removed.The morphology of burrows was related to sediment characteristics. Burrows were deeper, longer and more voluminous when sediment thickness was high. The volume of burrows decreased with increasing sediment torque. These results suggest that the morphology of burrows is related to the space available and the ease with which sediment it can be excavated. However, an important amount of variability remained unexplained, suggesting the presence of additional environmental variables or behavioural plasticity not considered by this study. Together, these results demonstrate that the spatial heterogeneity in the environmental factors can be translated to a spatial heterogeneity in the distribution of fiddler crabs.     

Endogenous rhythms and entrainment cues of larval activity in the horseshoe crab Limulus polyphemus

The American horseshoe crab, Limulus polyphemus (Linnaeus), typically inhabits estuaries and coastal areas with pronounced semi-diurnal and diurnal tides that are used to synchronize the timing of spawning, larval hatching, and emergence. Horseshoe crabs spawn in the intertidal zone of sandy beaches and larval emergence occurs when the larvae exit the sediments and enter the plankton. However, L. polyphemus populations also occur in areas that lack significant tidal changes and associated synchronization cues. Endogenous activity rhythms that match predictable environmental cycles may enable larval horseshoe crabs to time swimming activity to prevent stranding on the beach. To determine if L. polyphemus larvae possess a circatidal rhythm in vertical swimming, larvae collected from beach nests and the plankton were placed under constant conditions and their activity monitored for 72 h. Time-series analyses of the activity records revealed a circatidal rhythm with a free-running period of ≈ 12.5 h. Maximum swimming activity consistently occurred during the time of expected falling tides, which may serve to reduce the chance of larvae being stranded on the beach and aid in seaward transport by ebb currents (i.e., ebb-tide transport). To determine if agitation serves as the entrainment cue, larvae were shaken on a 12.4 h cycle to simulate conditions during high tide in areas with semi-diurnal tides. When placed under constant conditions, larval swimming increased near the expected times of agitation. Thus, endogenous rhythms of swimming activity of L. polyphemus larvae in both tidal and nontidal systems may help synchronize swimming activity with periods of high water and inundation.     

Behavioural evidence for body colour signaling in the fiddler crab Uca perplexa (Brachyura: Ocypodidae)

In the reproductive season, mature females of the fiddler crab Uca perplexa leave their burrows and wander about their habitat for mating. To clarify whether the fiddler crabs respond to colour or luminosity, I examined the behavioural responses of the males to the wandering females before and after the females were painted white, red, black or blue. The behaviours of the males were categorized into three types: lateral–circular wave and lateral–straight wave for courting, and repelling. Before painting, almost all of the males courted the females. After painting, significantly fewer males courted the red-, black- and blue-painted females than courted the white-painted females. These results mean that the fiddler crabs can discriminate colours or luminosity. The role of body colour as a visual signal in crab society is discussed.     

Approach of females to magnified reflections indicates that claw size of waving fiddler crabs correlates with signaling effectiveness

Male sand fiddler crabs, Uca pugilator, wave a claw to attract females to a breeding burrow. The effect of claw size on the likelihood of attracting mate-seeking females is little studied although in some other species females preferentially approach larger males. We used paired mirrors to reflect different sized images of the same male in a South Carolina (USA) back-beach habitat. Use of mirrors controlled for waving rate (but not velocity), waving motion, claw color, and claw shape. Female choice was attributed to instances in which a female contacted one of two mirrors. Paired mirrors were inclined toward one another in an arena defined by blinds and containing a single male. Two reflections of the male were visible to females moving approximately 50 cm toward the mirrors. The male was behind a small internal blind and not directly visible. In one-half of the trials, a non-magnifying mirror was placed at the bottom of mirrors so that only the elevated claw was magnified. Thus, body and burrow size and apparent distance were controlled. Receptive females preferred the larger reflection whether or not the body of the male was magnified, suggesting the importance of claw size. Non-receptive females did not exercise a choice. Control arenas, without a male, rarely attracted females. The results suggest that females choose on the basis of claw size. Selection on females may favor response to larger-clawed males because use of the claw in contests between males over burrows maintains the honesty of claw size as a signal of burrow quality.     

Endogenous rhythms of locomotion in the American horseshoe crab, Limulus polyphemus

American horseshoe crabs (Limulus polyphemus) exhibit clear circadian rhythms of visual sensitivity in the laboratory and in the field they exhibit seasonal patterns of mating behavior that are closely associated with the tides. Recent reports suggest that Limulus locomotor activity may be controlled by endogenous circadian and/or circatidal clocks and that light:dark (LD) cycles may affect the rhythmic output of both of these clocks. In this study, we examined locomotor behavior in the laboratory to determine the extent of this endogenous activity and to examine the influence of LD cycles on these rhythms. Thirty-three L. polyphemus were captured during the breeding season and their activity was monitored with activity boxes and “running wheels” in seawater kept at constant temperature and salinity. Activity patterns were analyzed using visual inspection of actograms and Chi-square and Lomb-Scargle periodograms. Overall, 36% of the animals was significantly more active during L, while only 12% was more active during D (52% showed no preference). Circatidal rhythms were observed in LD in 67% of the horseshoe crabs. Surprisingly, LD cycles appeared to synchronize these rhythms at times. In DD, the majority of animals tested (63%) exhibited circatidal rhythms that persisted for at least seven days. Overall, the results demonstrate that an endogenously controlled tidal rhythm of locomotion operates during, and significantly after, the breeding season in this species. In addition, the present results are consistent with the presence of circalunidian oscillators controlling these rhythms.     

Semilunar courtship rhythm of the fiddler crab<Emphasis Type="Italic"> Uca lactea</Emphasis> in a habitat with great tidal variation

Semilunar courtship rhythm is a widely distributed phenomenon among fiddler crabs in the genus Uca (Decapoda, Ocypodidae). Typically, synchronous courtship has been reported to peak near spring tides. To determine whether a region of large tidal variation shifts reproductive activity, we measured the frequency of specific courtship behaviors including claw-waving and semidome building for U. lactea males on Kanghwa Island, Korea. We found that synchronized courtship for U. lactea peaked near neap tides, whereas near the spring tides, seawater flooded the habitat and males predominantly fed on the mudflat. Although active females, which hold their burrows and usually feed on the mudflat, are abundant near to spring tides, males rarely claw-waved to attract females. This pattern is atypical for the species because other populations of U. lactea on Japan and Taiwan are synchronous around spring tides. We suggest that males invest most of their time in feeding during spring tides because foraging is limited during neap tides. During neap tides, males feed infrequently and thus expend stored energy on courtship signals. We conclude that patterns of reproductive synchrony may be dependent on food availability in periodically changing environments.     

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.