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.     

Predation by red-jointed fiddler crabs on congeners: interaction between body size and positive allometry of the sexually selected claw

The enlarged (major) claw of male fiddler crabs is used in contestsover breeding burrows and is waved to attract females. We recentlydiscovered that males of the red-jointed fiddler crab, Uca minax,also use the claw to kill smaller-sized fiddler crabs, U. pugnaxand U. pugilator, with which they co-occur in Atlantic coastsalt marshes. Large U. minax males use walking legs or the enlargedclaw to capture prey feeding on moist sand flats. On sand flats,small U. minax males and females are much less common than largemales, suggesting that large males move onto sand flats to seekprey. Males of prey species use the major claw against attackingpredators and, consequently, are more likely than females toescape. In laboratory experiments, large U. minax males weremore likely to attack and kill small-clawed males and femalesthan large-clawed males, consistent with a preference for morevulnerable, less threatening prey. The size of the major clawis a positive allometric function of body size. The allometricfunction varies little among species. Also, the mechanical advantageand indices of closing speed and closing force of the majorclaw, when corrected for body size, are not consistently greaterin U. minax relative to prey species. Thus, predation by U.minax males may reflect the opportunity afforded by larger bodysize and positive allometric growth, which result in a majorclaw that is more massive than the prey it is directed against.     

Acclimation of Intertidal Crabs

SVXOPSIS. Intertidal crabs are subjected to marked fluctuationsin environmental factors. Temperature and salinity influencethe distribution of organisms both on a latitudinal basis andalong a subtidal to terrestrial gradient. These factors areoperative on all stages of the life history. Resistance-adaptations. Adult fiddler crabs (genus Uca) fromthe temperate zone are more resistant to low temperature thantropical species. Also, the tolerance to low temperature ofcrabs from the temperate zone is greatly influenced by thermalacclimation, i.e. cold-acclimated crabs are more resistant thanwarm-acclimated animals. In contrast, tropical species havelimited adaptive ability. At elevated temperatures no consistentdifference in the lethal limits of crabs from tropical and temperatezones is observed. In contrast with the adults, larvae of tropicalspecies are cold-resistant. However, the larvae of all speciestested are more sensitive to reduced salinities than are theadults. Capacity-adaptatioyis. Subtidal species of crabs tend to havea lower level of respiratory performance, as measured by variousindices, than crabs from the intertidal zone. The metabolicresponse of fiddler crabs from the temperate zone is more labileat low temperature than in species from the tropical zone. Atelevated temperature the reverse response is observed. On aninterspecific basis, differences in the metabolictemperaturepatterns of acclimation of latitudinally separated populationsof U. pugilator are observed when based on the hypotheticalschemes of Bullock, Precht, and Prosser.     

Visual and Acoustical Signalling during Courtship by Fiddler Crabs (Genus Uca)

Male fiddler crabs (Genus Uca) employ both visual and acousticalsignals to attract females for mating. In U. pugilator and severalother American species, the males attract females during theday first by waving, then by producing sounds just within theirburrows. At night, the males produce sounds at low rates, butwhen touched by a female, they increase their rate of soundproduction. In the European species, U. tangeri, many elements of courtshipare similar to those in U. pugilator, but two types of soundsare produced. One of these, the short drumwhirl, appears tosubstitute for waving when the male is temporarily obscuredfrom the female during his diurnal courtship activities. Thelong drumwhirl is used under different circumstances. The acoustical responses of a male to a female influence thecourtship behavior of other males in the area. When sounds fromstimulated males are played back to test males during the day,their lates of waving increase. At night, the playbacks elicitincreases in rates of sound production. The influence of tidal oscillations, temperature, and lightcycles on the behavior of males is discussed. Courtship activities of aquatic crabs are compared to thoseof terrestrial Brachyura. In aquatic forms, courtship may beabsent or, if present, does not involve elaborate signallingby the male. Chemical or visual cues at close range are themost important stimuli. In several genera of terrestrial crabs,visual signalling for prolonged periods is common, and soundsare often emitted by males to "call" females from their burrowsto the surface for mating. Some of the factors that may accountfor differences in courtship activities in aquatic and terrestrialspecies are discussed.     

Display Specificity and Reproductive Isolation in the Fiddler Crabs,Uca panacea and U. pugilator

The factors responsible for reproductive isolation between two sibling species of fiddler crabs (Uca pugilator and U. panacea) were studied under both laboratory and field conditions. ♂♂ showed species differences in their visual and acoustical displays. These differences were exaggerated in the overlap zone, where U. pugilator showed character displacement of its acoustic signals. ♀♀ of U. pugilator confined with ♂♂ of U. panacea produced fewer clutches of young. Occasionally, forced matings took place in the laboratory, resulting in hybrids suffering greater mortality through development. The data indicate that both premating (behavioral) as well as postmating (higher larval mortality) barriers act to prevent interbreeding.     

Some Thermal Requirements of Fiddler Crabs of the Temperate and Tropical Zones and Their Influence on Geographic Distribution

Thermal stress is one environmental parameter that has greatlyinfluenced the migration of crustaceans from the sea to land.Since a greater number of species of terrestrial crabs are foundin the tropics than in the temperate zone, comparative studiesof the influence of temperature on latitudinally separated populationswere undertaken. Two tropical species, U. rapax and U. thayeri,may occur as far north as St. Augustine, Florida, or, followinga severe winter, may be rare north of Cape Kennedy. The lethaleffect of the low temperatures recorded during one severe winter(1957–58) is supported by laboratory studies in whichLD₅₀, deaths occurred in 4.5 days at 10°C for U. rapax acclimatedto 18°C. The experiment demonstrates that U. rapax cannotacclimate to and survive low temperatures. This contrasts markedlywith the situation in semi-terrestrial crabs of the temperatezone, which are able to acclimate to cold. The distribution of Uca around Cape Cod Bay correlates wellwith the coastal hydrographic thermal gradient and supportsPassano's suggestion that temperatures below 20° may belimiting as they inhibit proecdysis in U. pugnax. Such an inhibitionis found experimentally in U. pugilator and in the tropicalspecies, U. rapax. It is hypothesized that a shift in the thermodynamicsof the processes underlying molting has not occurred in Ucaof the temperate zone. The paucity of semi-terrestrial Brachyurain the temperate zone may be due to the failure of many speciesto evolve capacity-adaptations to carry out all requisite lifeprocesses at temperatures below 20°, or the resistance-adaptationsnecessary to survive the low temperatures of winter.     

Growth,survival, and metal content of marsh invertebrates fed diets of detritus from Spartina alterniflora Loisel. and Phragmites australis Cav. Trin. ex Steud. from metal-contaminated and clean sites

Marsh vegetation plays an important role in trophic ecology of estuaries. Once broken down to detritus, it is an important food source for manyorganisms. In Atlantic Coast marshes, the reed Phragmites australis hasbeen invading many areas once dominated by smooth cordgrass, Spartina alterniflora. In this study we evaluated the growth of and trophictransfer of metals to estuarine invertebrates when fed diets of detritus fromthese different plant species. Decaying leaves from populations of Phragmites, natural Spartina, and restored Spartina from boththe Hackensack Meadowlands, New Jersey, and the more pristineAccabonac Harbor of East Hampton, New York, were collected from themarsh surface in the spring. Decaying leaves were pureed and fed to thefiddler crabs Uca pugnax and U. pugilator, and to the grassshrimp Palaemonetes pugio. In fiddler crabs we monitored limbregeneration, molting and weight. U. pugilator regenerated limbs andmolted equally well on all six diets. Most of the U. pugnax arrestedgrowth midway through regeneration on all 6 diets. A repeat experimentwith smaller crabs, which did complete the process, found no consistentdifferences among the six diets and control food, although control food andPhragmites detritus had higher N concentrations than the Spartinadetritus. Grass shrimp fed all six diets did not survive beyond 3 weeks. Inanother experiment using HM sediments from each vegetation type(containing detritus, meiofauna, and microflora), survival was equally highamong treatments and the shrimp fed sediments from the restored Spartina site or control food grew better than those fed sediments fromthe Phragmites or natural Spartina sites. Although metalconcentrations in detritus varied between sites and plant species, the crabsof each group did not differ in metal concentrations after the feedingexperiment. Our data do not support the general assumption that Phragmites leaf detritus is of poorer nutritional quality than Spartinaalterniflora leaf detritus to estuarine consumers.     

Regulation of sex-specific feeding behavior in fiddler crabs: physiological properties of chemoreceptor neurons in claws and legs of males and females

This study examined properties of chemoreceptor neurons in the claws and legs of the fiddler crabs Uca pugilator and U. pugnax. The primary goal was to establish the neural basis of previously observed greater female sensitivity to feeding stimulants, and secondarily to compare physiological properties of chemoreceptor neurons in these semi-terrestrial crustaceans with those of fully aquatic forms. Sensitivity of chemoreceptor neurons in claws and legs is sex-specific; individual neurons of females respond to lower stimulus concentrations than male chemoreceptor neurons, and equivalent concentrations elicit greater spiking in female vs male chemoreceptor neurons. Thus, the population of chemoreceptor neurons in females expresses lower thresholds and greater average sensitivity than in males. Greater sensitivity of claw neurons explains observations indicating that females continue to feed at food levels too low to stimulate males. Sensitivity differences in leg neurons of males vs females have no clear behavioral correlate, but suggest that females can orient to more dilute stimuli than males. Chemoreceptor neurons of fiddler crabs have low sensitivities and slow rates of adaptation compared to other crustaceans. Also, neurons in claws adapt less slowly than neurons in legs, which may reflect subtle differences in the chemical stimulus environment experienced by claws vs legs.     

Burrowing,root mat density,and the distribution of fiddler crabs in the eastern United States

This paper discusses the distribution of two species of fiddler crabs across an estuary. The possibility that sediment size, physical factors (or micro-climate), or food limitation could lead to the observed distributions is discussed, and evidence is presented to show that these factors probably play a minor rôle in controlling the distribution of Uca pugnax (Smith) and U. minax (Le Conte) across an estuary. The results show that the best correlate of the dominant species of Uca in a particular habitat is the density of the root mat of that area. A series of experiments showed that variations in the abundance of large U. pugnax at two tide levels in a marsh very closely parallel the ability of U. pugnax to burrow in substrata taken from those same two areas of the marsh. These two areas have significant differences in density of the root mat.     

Independent and interactive effects of two facilitators on their habitat‐providing host plant,Spartina alterniflora

The role of habitat‐providing species in facilitating associated species abundance and diversity is recognized as a key structuring force in many ecosystems. Reciprocal facilitation by associates, often involving multiple species, can be important for the maintenance of the host species. As with other multi‐species interactions (e.g. multiple predator effects), non‐additive relationships may be common among these associates, yet relatively few studies have examined potential interactions among multiple facilitator species. We combined field surveys and a mesocosm experiment to examine the independent and interactive effects of two co‐occurring facilitator species, ribbed mussels Geukensia demissa and fiddler crabs Uca pugilator, on their host salt marsh plant species, cordgrass Spartina alterniflora. We also experimentally examined how these relationships varied across different host plant genotypes. Overall, facilitator effects increased with increasing facilitator density. There was a significant interaction between mussel and fiddler crab presence, indicating that the effects of each species on cordgrass were dependent on the presence of the other facilitator species. In addition, there were strong interactions among mussels, fiddler crabs, and plant genotype, with greater variation in the performance of individual genotypes when fiddler crabs were absent. Our work reinforces the importance of considering multiple responses when assessing the functional redundancy of co‐occurring facilitators, as species are seldom completely redundant across the range of services they provide. It also highlights that the strength and direction of species interactions can vary due to genetic variation within the interacting species.     

Effect of cadmium exposure on lactate dehydrogenase activity in the hepatopancreas and abdominal muscle of the fiddler crab,Uca pugilator

1. Lactate dehydrogenase (LDH) activity in the hepatopancreas and abdominal muscle of fiddler crabs, Uca pugilator, was determined after 24 and 48 hr of exposure to 2 ppm cadmium chloride.2. For the cadmium exposed crabs, LDH activity in the hepatopancreas decreased, whereas that in the abdominal muscle increased.3. The increased LDH activity in the abdominal muscle may reflect increased dependence on anaerobic carbohydrate metabolism in fiddler crabs exposed to cadmium in their environment.     

Osmoregulation by six species of fiddler crabs (Uca) from the Mississippi delta area in the northern Gulf of Mexico

Six species of fiddler crabs (Ocypodidae: Uca) were collected for osmoregulation studies from 25 locations near the delta of the Mississippi River in the northern Gulf of Mexico. Three of the species are classified as members of the Celuca subgenus, Uca spinicarpa, Uca panacea and Uca pugilator, while the remaining three are in the Minuca subgenus, Uca minax, Uca longisignalis and Uca rapax. In the field, U. minax, U. spinicarpa and, occasionally, U. longisignalis are found in freshwater habitats (FW; 0-299 mosM). Two Minuca species, U. longisignalis and U. rapax, are typically collected in brackish water habitats (BW; 300-629 mosM). On the other hand, U. panacea and U. pugilator are most abundant in eurysaline habitats (EH; >630 mosM). In the laboratory, populations of each species were challenged with media ranging from 30 to 3450 mosM (1-110‰). The FW species, U. spinicarpa and U. minax, did not tolerate osmotic concentrations >2100 mosM. The EH species, U. panacea and U. pugilator, however, tolerate concentrations >2800 mosM. The BW species, U. longisignalis and U. rapax, succumb to osmolalities between 2100 and 2800 mosM. Each species keeps its hemolymph concentration fairly constant in 30-1400 mosM solutions. The [ISO], isosmotic medium concentration (in mosM), is calculated for each taxon: U. minax, 659; U. spinicarpa, 682; U. longisignalis, 693; U. rapax, 769; U. pugilator, 816; and U. panacea, 822. In media with >1600 mosM, each species expresses different osmoregulating capabilities. The FW species, U. spinicarpa and U. minax, cannot control hemolymph osmolality above 1500 mosM while the BW-EH species, U. panacea, U. pugilator and U. rapax, regulate hemolymph values in media up to 2300 mosM. Within the FW/BW species U. longisignalis, the ability to osmoregulate corresponds with site of collection. Specimens from FW populations do not regulate as well as those from BW if challenged with hypertonic media. If adapted to a 1800 mosM in the laboratory, survivorship for U. longisignalis shifts to the right and the [ISO] increases to 832 mosM. This suggests that this species adapts to acute hypertonic conditions by tolerating elevated internal osmolality. Generally, these observations extend our knowledge about the physiological capabilities of fiddler crabs from different salinity populations across the northern Gulf of Mexico.     

Distal Retinal Pigment of the Fiddler Crab,Uca pugilator: Release of the Dark-Adapting Hormone by Methionine Enkephalin and FMRFamide

The neuropeptides methionine enkephalin and FMRFamide, when injected into intact fiddler crabs, Uca pugilator, produce dark adaptation of the distal retinal pigment. Furthermore, both neuropeptides stimulate release of distal retinal pigment dark-adapting hormone activity from the isolated eyestalk neuroendocrine complex. It is hypothesized that both neuropeptides, when injected into intact fiddler crabs, act only indirectly on the distal retinal pigment, by stimulating release of this dark-adapting hormone.     

Comparative Studies of Reproductive Behavior in Mantis Shrimps and Fiddler Crabs

Comparative ethological studies of closely related species canplace into clear perspective the functions of behavioral traitsas species diverge and adapt to different environments. In thispaper we contrast the reproductive behavior of species withintwo crustacean groups: the mantis shrimps (stomatopods) andthe fiddler crabs (genus Uca). For the stomatopods, we identifyprobable selective relationships between features of species'ecology and their mating systems. Population density, the kindand availability of shelters in which these shrimps live, theintensity of predation and spawning cycles all play importantroles in molding reproductive behavior in this group. Associationsbetween the ecology and mating systems of fiddler crabs havebeen discussed recently elsewhere. Here we focus on a comparativeanalysis of sexual communication in these crabs. A study ofcompetitive courtship signaling and mate choice in the fiddlercrab Uca beebei has shown that certain male courtship signalsare highly attractive probably because they exploit female sensory-responsesystems that have been molded by selection for escape from predators.Interspecific comparison of male courtship displays and theresponses of females to these displays suggest that sensoryexploitation may play an important role in the evolution ofsexual signals in the genus. Comparative studies have advancedour understanding of how natural and sexual selection affectthe reproductive behavior of both stomatopods and fiddler crabs.     

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.     

Risk-taking behavior predicts aggression and mating success in a fiddler crab

Evidence is growing that an individual's propensity to takerisks in the presence of a predator is correlated to behaviorsthat can affect individual fitness. We examined whether risk-takingbehavior predicts aggression, surface activity levels, and matingsuccess in male fiddler crabs, Uca mjoebergi. Risk-taking behaviorwas highly consistent among individuals, remained stable overtime, and was unrelated to male size. We found that males thattook greater risks in the presence of a potential predator alsobehaved more aggressively when searching for a new territory.In addition, bold males exhibited higher surface activity levelsand spent more time courting females compared with their shycounterparts. Although risk-taking behavior was independentof other sexually selected traits, it accurately predicted malemating success in U. mjoebergi. We suggest nonsexually selectedtraits, such as risk taking, may represent important behavioralpredictors of success in other species.     

Effect of Eyestalk-Ablation on Circulating Ecdysteroids in Hemolymph of Snow Crabs, Chionoecetes opilio: Physiological Evidence for a Terminal Molt

Bering Sea snow crabs (Chionoecetes opilio) are a commerciallyimportant crab harvested in the Bering Sea. Optimal managementof this species requires an understanding of the biology ofthis crab that is currently incomplete. Fisheries managers applya continuous growth model in their management of snow crab,which assumes that male crabs increase in size throughout theirlifespan. Male snow crabs undergo a morphometric molt that leadsto a disproportionate increase in chelae size and it is stilldebated whether this molt is associated with a terminal molt.This study was conducted to determine whether adult male C.opilio are anecdysic. Using current knowledge of the hormonalregulation of crustacean growth, snow crab physiology was manipulatedto induce an increase in molting hormones (ecdysteroids). Sincefemale snow crabs are known to undergo a terminal molt afterattaining reproductive maturity, we compared ecdysteroid levelsin eyestalk-ablated terminally molted females, small-clawedmales and large-clawed males. Snow crabs were collected fromthe Bering Sea and maintained in circulating seawater at approximately6°C. Animals were either eyestalk-ablated or left intact.Ecdysteroid levels in hemolymph were quantified using an enzyme-linkedimmunosorbant assay (ELISA). Circulating ecdysteroids were significantlyhigher in small-clawed male crabs when compared to large-clawedmales or terminally molted females. Eyestalk-ablation increasedcirculating ecdysteroids in small-clawed males, but had no significanteffect on circulating ecdysteroids in large-clawed males orin terminally molted females.     

Morphological diversity of setae on the second maxilliped of fiddler crabs (Decapoda: Ocypodidae) from the southwestern Atlantic coast

Fiddler crabs are deposit feeders, and use the setae on their mouth appendages to manipulate sediment particles to extract food. The number of spoon‐tipped setae on the second maxilliped is frequently related to the distribution of fiddler crabs on estuarine sediments, but no study has compared the morphological diversity of these setae among multiple fiddler crab species. Here, we describe and classify the setae of the second maxillipeds of the nine Uca spp. known from the Brazilian coast. The second maxilliped of each species was examined by scanning electron microscopy. Six types of setae (five papposerrate, and one pappose) were described on the meropodite of the second maxilliped. Among the papposerrate setae, one type had a spoon‐like tip, and the morphology of this type, especially the degree of curvature, differed between species. Members of Uca leptodactylus, U. uruguayensis, and U. maracoani had highly concave spoon‐tipped setae. In U. rapax and U. cumulanta, the setal tip was moderately curved, while in U. thayeri, U. burgersi, and U. mordax, this curvature was slight. At the other extreme, the meropodite of the second maxilliped of U. vocator lacked setae altogether. This is the first study that describes differences in the degree of curvature of spoon‐tipped setae in fiddler crabs. This trait may be strongly related to the distribution of these fiddler crabs on different estuarine substrates.     

The fiddler crab,Minuca pugnax,follows Bergmann's rule

Bergmann's rule predicts that organisms at higher latitudes are larger than ones at lower latitudes. Here, we examine the body size pattern of the Atlantic marsh fiddler crab, Minuca pugnax (formerly Uca pugnax), from salt marshes on the east coast of the United States across 12 degrees of latitude. We found that M. pugnax followed Bergmann's rule and that, on average, crab carapace width increased by 0.5 mm per degree of latitude. Minuca pugnax body size also followed the temperature–size rule with body size inversely related to mean water temperature. Because an organism's size influences its impact on an ecosystem, and M. pugnax is an ecosystem engineer that affects marsh functioning, the larger crabs at higher latitudes may have greater per‐capita impacts on salt marshes than the smaller crabs at lower latitudes.