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.     

Shell use by juvenile fiddler crabs Uca pugnax and U. pugilator

The present study documents for the first time shell use by juvenile fiddler crabs in the salt marsh. Twenty visits were made to six salt marsh sites at Tybee Island, Georgia between 2007 and 2009. One hundred empty Littorina irrorata shells were collected at each site on each field trip. Juvenile carapace width was measured, crabs sexed, and species identification completed using RFLP analysis. Shell use of up to 79% was observed. Two species of fiddler crabs were found in empty shells, Uca pugnax and U. pugilator. U. pugnax was the dominant species at all sites representing 62-84% of the juvenile fiddler crab population. Juvenile sex ratios were female-biased (1.7:1) at all six sites. Juvenile size did not vary significantly between species but males of both species were significantly larger than females. Size frequency distribution of carapace width revealed that shell use varied with size and sex. In the 3 to 4 mm size class, juvenile females outnumbered juvenile males in empty L. irrorata shells while in the 5 to 6 mm size class and greater, juvenile males outnumbered juvenile females in shells. Significantly more juvenile fiddler crabs were found in empty shells during flood than ebb tide at 3 of the sites. This discovery illuminates the resourcefulness of juvenile fiddler crabs and provides another mechanism that might enhance survival.     

Metabolic responses of sand fiddler crabs, Uca pugilator, in northwest Florida to seasonal temperature change

Metabolic responses of sand fiddler crab, Uca pugilator, populations in northwest Florida are greatly influenced by seasonal temperature fluctuations. Crabs acclimated at 20 °C and immediately transferred to either 14 or 26 °C produced an acute metabolic response with respective temperature quotient (Q₁₀) values of 3.46 and 3.91. Crabs acclimated at 10 and 20 °C exhibited a Q₁₀ of 2.62 indicating a partial compensation response. A brumation (reverse) response (Q₁₀ value of 20.11) was observed for acclimated crabs between 5 and 10 °C. Brumation is advantageous during winter when food supplies are scarce and crabs must survive extensive periods of inactivity.     

Experimental infections of Orchitophrya stellarum (Scuticociliata) in American blue crabs (Callinectes sapidus) and fiddler crabs (Uca minax)

Outbreaks of an unidentified ciliate have occurred on several occasions in blue crabs from Chesapeake Bay held during winter months in flow-through systems. The parasite was initially thought to be Mesanophrys chesapeakensis, but molecular analysis identified it as Orchitophyra stellarum, a facultative parasite of sea stars (Asteroidea). We investigated the host-parasite association of O. stellarum in the blue crab host. Crabs were inoculated with the ciliate, or they were held in bath exposures after experimentally induced autotomy of limbs in order to determine potential mechanisms for infection. Crabs inoculated with the ciliate, or exposed to it after experimental autotomy, rapidly developed fatal infections. Crabs that were not experimentally injured, but were exposed to the ciliate, rarely developed infections; thus, indicating that the parasite requires a wound or break in the cuticle as a portal of entry. For comparative purposes, fiddler crabs, Uca minax, were inoculated with the ciliate in a dose-titration experiment. Low doses of the ciliate (10 per crab) were sometimes able to establish infections, but high intensity infections developed quickly at doses over 500 ciliates per crab. Chemotaxis studies were initiated to determine if the ciliate preferentially selected blue crab serum (BCS) over other nutrient sources. Cultures grown on medium with BCS or fetal bovine serum showed some conditioning in their selection for different media, but the outcome in choice experiments indicated that the ciliate was attracted to BCS and not seawater. Our findings indicate that O. stellarum is a facultative parasite of blue crabs. It can cause infections in exposed crabs at 10–15 °C, but it requires a portal of entry for successful host invasion, and it may find injured hosts using chemotaxis.     

Semi-monthly reproductive cycles in male and female fiddler crabs, Uca pugnax

In the fiddler crab, Uca pugnax, numbers of displaying males, numbers of burrows with half-dome superstructures (hoods), numbers of females with decalcified vulvar opercula, and numbers of females hatching their eggs, all follow a semi-monthly cycle with peaks near the time of the spring tides. These are all aspects of reproduction in Uca. Decalcification of female opercula, although necessary for mating and egg deposition, is neither necessary nor sufficient for behavioural response to courting males. In fact, behavioural responsiveness to males probably precedes decalcification in most cases. The different aspects of the reproductive cycle are coordinated so that females and males are ready to mate at about the same time near one spring tide, and the resulting eggs will be ready to hatch on the next spring tide. Two decalcified females which were kept isolated from males produced fertile clutches, suggesting that stored sperm from a previous mating may fertilize eggs in a later cycle. This may be one reason for the relative rarity of observed matings or precopulatory behaviour in Uca.     

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.     

A calibration method to generate seasonal hatching profiles for the fiddler crabs Uca pugnax (Smith, 1870) and Uca minax (LeConte, 1855) (Decapoda : Brachyura)

A statistical method for generating seasonal egg hatching profiles is applied to the brachyuran crabs Uca pugnax (Smith) and U. minax (LeConte) in New Jersey. Calibration experiments quantified the time course of egg development, using serial egg sampling at 1- to 2- day intervals from marked ovigerous females maintained in cages in the field. Egg stage was ranked from 1 to 10, based on morphological changes during development. Equations for predicting the number of days remaining until hatching from egg developmental stage were obtained from the calibration experiments, using stepwise polynomial regression. To cover the reproductive season, three consecutive calibration experiments using 15 or more females were run for U. pugnax; two for U. minax. Significant seasonal differences in the time course of egg development were detected. Weekly collections of females for each species were made; the date of larval release for each ovigerous female was predicted from the proximate calibration equation, yielding weekly hatching profiles. Weekly hatching profiles were summed to obtain seasonal hatching profiles. The average number of broods produced per female over the reproductive season was 1.9 for U. pugnax and 1.6 for U. minax. Hatching peaks for both species were associated with spring tides. The merits of this and other methods used to estimate daily variation in egg hatching of crabs are discussed.     

Carapace color change in Uca pugilator as a response to temperature

Brachyurid species have various coloration patterns that fluctuate among individuals due to behavioral interactions or a changing environment. In the fiddler crab Uca pugilator, the historical notion is that color change is regulated by circadian rhythms. In this study, color patterns of U. pugilator were affected by changes in temperature; crab carapaces become light in warm temperatures and dark in colder temperatures. The response to temperature occurred within 15 min (often within 5 min of exposure to the treatment), suggesting that temperature may play a key role in color change, and it is activated not through a hormonal pathway, but rather a direct response of the chromatophores. Color change occurred differently between day and night for each sex, which implies that endogenous rhythms are not the sole mechanism for color change in U. pugilator. Both sexes became darker in the cold treatment, however there were differences between the sexes with high temperatures. Females did not show significant coloration changes, while males became significantly lighter and more speckled. Results from this study suggest sex-specific constraints in response to temperature that could be associated to morphological body mass to surface area ratios, as males have large cheliped, which may be involved in thermoregulation. Constraints could also be associated to different diel activities that each sex performs or visual cues associated with mating behaviors.     

Stereotypy and the effects of temperature on some spatio-temporal subcomponents of the ‘courtship wave’ in the fiddler crabs Uca minax (Le Conte) and Uca pugnax (Smith) (Brachyura,Ocypodidae)

The effects of air temperature on some spatial and temporal subcomponents of the male wave display were investigated under laboratory conditions in the fiddler crabs, Uca minax and Uca pugnax. Also investigated were the stereotypies of these subcomponents. In general, wave duration, ascending and descending wave times decreased significantly with increasing air temperatures between 13 and 26 C. These decreases were generally not significant at air temperatures greater than 26 to 30 C. Wave frequency and the number of cheliped jerks (U. minax) increased with temperature, whereas the interwave time showed no systematic change with temperature. Uca pugnax had waves with discrete cheliped jerks at temperatures less than 20 C, whereas the wave appeared relatively smooth at higher temperatures. Temperature effects on waving varied between species and among behavioural contexts. Stereotypies of the wave subcomponents, as measured by the coefficient of variation, underwent no systematic change with temperature. However, stereotypies were different within and between species. Behavioural context also had an effect on subcomponent stereotypy. Comparisons of stereotypies yielded predictions concerning communication functions of waving subcomponents.     

Delayed courtship in the fiddler crab Uca musica terpsichores

Courting male fiddler crabs of the species Uca musica terpsichores congregate in the upper central portion of the colony, while receptive females leave their burrows located at the colony's periphery and wander among the communally displaying males prior to choosing a mate. I observed that courting males in a newly-established population were significantly smaller than courting males in large high-density colonies. This observation led to a series of translocation experiments designed to ascertain whether high population density influences the size (=age) at which males begin to court. Smaller courting males from a low-density population failed to court after being placed among larger courting males in a high-density population. The reciprocal translocation revealed that smaller noncourting males from the high-density population would start courting shortly after being placed in a low-density population. Smaller males placed in the high-density population were subsequently observed significantly further away from where they were initially placed than were larger males similarly translocated. The results suggest that smaller males delay courtship activities once they are forced, via encounters with larger males, to the periphery of the colony. I believe that both intrasexual selection (competition from larger males) and intersexual selection (female choice of large males) are responsible for the delay in male courtship activities.     

Habituation to disturbance in the fiddler crab (Uca annulipes) in its natural environment

Crabs of both sexes and of various sizes were subjected to a standardized test of ten consecutive disturbances in their natural habitat. Males and females reacted alike: 53 per cent became fully habituated within less than ten disturbances, and a further 12 per cent showed slower habituation. The remainder of the test individuals showed various other patterns of reactions. Reaction pattern and efficiency of habituation are size- (or age-) dependent, and they are modifiable by social interaction and by tide-determined activity phases. The structure and species composition (with regard to ocipodid crabs only) of the Uca habitat is considered briefly, and some observations on display and ‘rapping’ or drumming behaviour are reported.     

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.     

Developmental shift in the selective tidal-stream transport behavior of larvae of the fiddler crab Uca minax (LeConte)

Following hatching, larvae of the fiddler crab Uca minax (La Conte) are exported from the adult habitat in estuaries to coastal and shelf waters where they undergo development prior to re-entering estuaries as postlarvae (megalopae). Studies of the spatial distribution of both newly hatched zoeae (Stage I) and megalopae indicate they undergo rhythmic vertical migrations associated with the tides for dispersal and unidirectional transport (selective tidal-stream transport) both within estuaries and between estuaries and the nearshore coastal ocean. We tested the hypothesis that U. minax zoeae possess a circatidal rhythm in vertical migration that facilitates offshore transport in ebb tidal flows, while postlarvae (megalopae) return to estuaries using a similar flood-phased endogenous rhythm. We also determined if the expression of the rhythm was influenced by the salinity conditions zoeae and megalopae experience as they transition between low-salinity regions of estuaries and high-salinity coastal waters. Stage I zoeae were collected by holding ovigerous female crabs in the lab until hatching. Megalopae were collected from the plankton and identified to species using molecular techniques (PCR-RFLP). Under constant laboratory conditions, both zoeae and megalopae exhibited endogenous circatidal rhythms in swimming that matched the principal harmonic constituent of the local tides (12.39 ± 0.07 h; X¯ ± SE). Upward swimming in Stage I zoeae occurred 2.5-4 h after high tide near the time of expected maximum ebb currents in the field. Rhythmic swimming of megalopae occurred slightly earlier in the tide (2.5 ± 0.09 h after high tide; X¯ ± SE) but was not entirely synchronized with flood currents, as expected. Salinity conditions had no apparent effect on the expression or pattern of the rhythms. Results indicate that this circatidal rhythm forms the behavioral basis of selective tidal-stream transport (STST) in early stage U. minax zoeae, but does not undergo a sufficient phase shift to account for vertical distribution patterns exhibited by megalopae in the field.     

Male size and reproductive success in the communal courtship system of the fiddler crab Uca rapax

Reproductive success of male fiddler crabs Uca rapax depends on the numbers of females a male can attract, and the fecundity of these females. The latter correlates directly with fenale size. The size of a female in a mated pair is directly correlated with the size of the male. Therefore, older larger males experience greater reproductive success than smaller younger males. Females appear to be highly selective in mating, while males are not. The question of why females should select a male whose size is correlated with hers cannot be answered from the present study.