Colouration and colour changes of the fiddler crab, Uca capricornis: a descriptive study

Colour changes in animals may be triggered by a variety of social and environmental factors and may occur over a matter of seconds or months. Crustaceans, like fiddler crabs (genus Uca), are particularly adept at changing their colour and have been the focus of numerous studies. However, few of these studies have attempted to quantitatively describe the individual variation in colour and pattern or their adaptive significance. This paper quantitatively describes the colour patterns of the fiddler crab Uca capricornis and their ability to change on a socially significant timescale. The most dramatic changes in colour pattern are associated with moulting. These ontogenetic changes result in a general reduction of the colour pattern with increasing size, although females are more colourful and variable than similarly-sized males. Uca capricornis are also capable of rapid colour changes in response to stress, but show no endogenous rhythms associated with the semilunar and tidal cycles commonly reported in other fiddler crabs. The extreme colour polymorphism and the relative stability of the colour patterns in Uca capricornis are consistent with their use in visually mediated mate recognition.     

The fiddler crab Uca mjoebergi uses colour vision in mate choice

Although the role of colour in mate choice is well known, few tests of colour vision have been based on mating behaviour. Females of the fiddler crab Uca mjoebergi have recently been shown to use claw coloration to recognize conspecific males. In this study I demonstrate that the females use colour vision for this task; preferentially approaching yellow claws over grey claws regardless of their intensity while failing to discriminate between yellow claws differing in intensity. This is one of only a handful of studies confirming the involvement of colour vision in mate choice and the first conclusive evidence in fiddler crabs.     

Behavioral patterns of two fiddler crab species Uca rapax and Uca tangeri in a seminatural mangrove system

Zoos increasingly transform their exhibitions from traditional one‐species enclosures to more natural exhibits, that is, environments that capture part of an ecosystem including a selection of animals and plants that occur there. Thus, enhancing the experience of its human visitors while also allowing its residents to possibly show more natural behavior. In 2017 Royal Burger's Zoo (Arnhem, The Netherlands) created and opened a mangrove‐like environment containing fiddler crabs. Fiddler crabs display a broad range of behaviors, and this research examines which wild‐type behavior and behavioral patterns can be observed on a seminatural mudflat. The behavior shown by Uca rapax and Uca tangeri on the mudflat was counted each hour between 07:00 and 17:00. An asymmetric tidal regime was present in the enclosure including two high water periods. Various known fiddler crab behaviors, including waving and combat, were observed but no copulation. A clear pattern in exposed crabs on the mudflat was found, with low numbers visible in the early morning and the highest numbers present in the early afternoon, while number of visitors did not have a significant effect on this pattern. Interestingly, the highest abundances were not observed around the ebbing tide (07:00–09:00), as observed in the wild, but somewhat later, possibly due to the asymmetric tidal scheme or the interaction of tidal and daily rhythms. This study shows that in captivity, fiddler crabs indeed show a range of natural behaviors which is linked to the tidal and possibly daily rhythm as well.     

Factors Affecting Mating Tactics in the Fiddler Crab, Uca vocans hesperiae

The mating strategies of male fiddler crabs are variable and highly flexible within species. In this study I examine three types of mating strategy used by individual male Uca vocans hesperaie. The most common strategy, termed a ‘standard gambit’, where males approached females at their burrow entrance and initiated courtship, accounted for 63% of mating attempts and 75% of successful matings. The rarest strategy (4% of mating attempts) was the ‘dig out’, where males attempted to mate with females whose burrows they had excavated. This strategy accounted for 19% of successful matings. ‘Herding’ behaviour which involved a male attempting to herd a female into a burrow and mate, contributed 33% of mating attempts but were generally unsuccessful, accounting for only 2.6% of successful matings. Males used more than one strategy during the study period. Smaller males used the standard gambit strategy more often than herding or dig outs while larger males used the herding strategy more often. There was no relationship between male size and mating success and males did not preferentially mate with females of a certain size. The predominant strategy adopted by males over the lunar cycle depended on female behaviour. Herding behaviour was induced by female wandering which escalated at full moon. Standard gambits were the commonest strategy adopted at and around new moon. The low success rate of male mating attempts (16%) indicates a reluctance by females to mate multiply. This may lead to conflict between the sexes because in fiddler crabs there is last male sperm precedence.     

Courtship herding in the fiddler crab <Emphasis Type="Italic">Uca elegans</Emphasis>

Male and female animals are not always complicit during reproduction, giving rise to coercion. One example of a system that is assumed to involve sexual coercion is the mate herding behaviour of fiddler crabs: males push females towards the home burrow with the goal of forcing copulation at the burrow entrance. We recorded and analysed in detail the courtship behaviour of a North Australian species of fiddler crab Uca elegans. Courtship was composed of four main phases: broadcast waving, outward run, herding and at burrow display. During interactions males produced claw-waving displays which were directed posteriorly towards the female and which varied in timing and structure depending on the courtship phase. We suggest that courtship herding in U. elegans is driven primarily by mate choice for the following reasons, (1) females can evade herding, (2) no other reproductive strategies were observed, (3) males broadcast their presence and accompany courtship with conspicuous claw waves, and (4) the behaviour ends with the female leading the male into the home burrow. As an alternative function for herding in U. elegans we suggest that the behaviour represents a form of courtship guiding, in which males direct complicit females to the correct home burrow.     

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.     

Evaporative water loss, corporal temperature and the distribution of sympatric fiddler crabs (Uca) from south Texas

Desiccation and thermal stress are among the primary factors limiting terrestriality in crustaceans. Water loss was estimated as weight change in five sympatric species of Uca from south Texas for periods up to 7 hr in dry air. Simultaneously, corporal temperature was measured with a thermocouple placed under the carapace. To estimate integumental permeability to water, 115 mm2 portions of dorsal carapace were glued to U-shaped tubes containing a crab Ringer's solution. These were exposed to dry air and water permeability was estimated from weight change. In whole-animal studies, most rapid weight loss occurred in the first 5 min of exposure to dry air as the body temperature fell below ambient (25 degrees C) in all species. The three most terrestrial species exhibited significant survival over more aquatic congeners after prolonged desiccation. The greatest rate of water loss was observed in Uca subcylindrica which lost 22.9+/-3.0% body weight. Uca panacea and Uca spinicarpa lost 14.1+/-1.6% and 18.5+/-1.8%, respectively. Based on blood osmolarity changes, Uca longisignalis and Uca rapax were more resistant to water loss than Uca subcylindrica under these conditions. Water loss from sections of the dorsal carapace were highest in Uca spinicarpa (10.4 mg/hr/cm2) and Uca longisignalis (8.9 mg/ hr/cm2). Uca subcylindrica and Uca panacea were intermediate (4.5 and 4.2 mg/hr/cm2) while Uca rapax expressed the lowest value (2.9 mg/hr/cm2). These observations support the notion that water loss can effectively lower body temperature in fiddler crabs. However, an inverse relationship between terrestriality and integumental permeability was not evident in these sympatric congeners. Ultimately a balance between physiological and behavioral mechanisms must be achieved for adaptation to the semi-arid habitats in south Texas.     

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.     

Microgeographic genetic structure of the fidller crab,Uca arcuata De Haan (Ocypodidae) in Taiwan

The genetic structure of Uca arcuata in Tanshui mangrove swamp of northern Taiwan was examined. Using as genetic markers, isozymes identified through starch gel electrophoresis indicate that there was moderate genetic differentiation among subpopulations within the population (FST = .085). Gene flow appeared high when estimated indirectly (Nm = 2.69). The results suggest that the patterns of genetic structure of Uca arcuata were influenced by the interaction of local selection due to microhabitat differences and gene flow among fiddler crab colonies in the mangrove swamp.     

Response of fiddler crabs (Uca tangeri) to video playback in the field

The fiddler crab Uca tangeri communicates using a visual waving display and a vibratory drumming signal, both thought to function in mate attraction. Using video playback techniques, images of an empty mudflat, a waving male, a threatening male, and a wandering female were presented to male subjects. All stimuli elicited similar levels of low-intensity waving, but significantly more high-intensity waves were elicited by the female and threatening male stimuli than the mudflat stimulus or the waving male stimulus. This concurs with other research that the waving display is used at a higher intensity to attract females. The threatening male stimulus failed to elicit the same response as an actual threatening male and we discuss the likely reasons for this. The subjects also spent a significantly greater proportion of time drumming during the male waving stimulus than during the female stimulus, suggesting that drumming functions in male-male competition as well as female attraction. Received: 18 October 1999 / Received in revised form: 25 November 1999 / Accepted: 10 December 1999     

Evidence for a two pigment visual system in the fiddler crab, Uca thayeri

Intraocular recordings were made from the eyestalks of dark-adapted fiddler crabs (Uca thayeri) during presentation of monochromatic light flashes of different wavelengths and intensities. Two types of signals were recorded in different experiments: slow potentials (electroretinogram) and fast potentials (spikes). The latter were also recorded in the presence of a continuous green or red adapting light. The resulting visual spectral-sensitivity curves, when fitted to rhodopsin-based visual pigment absorption spectra (from Dartnall nomograms), indicated the presence of two visual pigments, one with an absorption maximum near 430 nm, and the other with a peak absorption between 500 nm and 540 nm. The data also provided evidence for some differential bleaching of the pigments in the presence of a colored adapting light, but most of the adaptation effect was probably due to changes in screening pigment and neural desensitization or inhibition. These two observations suggest that an adequate substrate for color vision may exist in this and other species of fiddler crabs. The electroretinogram and spike-recording methods produced similar visual-sensitivity data, suggesting that latter technique, a much more efficient way of collecting data that is physiologically relevant, may be the method of choice for determining spectral sensitivity in crustaceans.     

Short-range Homing in Fiddler Crabs (Ocypodidae, Genus Uca): A Homing Mechanism not Based on Local Visual Landmarks

This field study investigated the spatial strategies and homing ability of two East African fiddler crabs, Uca lactea annulipes and Uca vocans hesperiae, using various experimental procedures. A clear-cut spatial segregation between the two species was observed, with U. l. annulipes occupying areas even a few centimetres above U. v. hesperiae . Females of both species were extremely faithful to their holes while the males exhibited behavioural differences. U. l. annulipes males were quite faithful to their holes; in contrast, U. v. hesperiae males, during a single low tide, visited several holes which they did not defend. Two displacement experiments were conducted in order to identify the stimuli that fiddler crabs use in their homing. In the 'dislocation experiment', the crabs were moved (by hand) 1 or 2 m away from their refuge and released. None of these crabs returned to its burrow and, moreover, the initial directions they followed were not homeward orientated. In the 'translation experiment', the crabs were passively translated while actively feeding. When the crabs attempted to go 'home' after the translation, they darted along a straight path that led them to points that would have corresponded to their homes if the translation had not taken place. After this first response crabs then commenced searching strategies that led half of them home. Short-range homing seems to depend strongly on information actively gathered during the outward journey, other than local visual landmarks.     

Reproducing on Time When Temperature Varies: Shifts in the Timing of Courtship by Fiddler Crabs

Many species reproduce when conditions are most favorable for the survival of young. Numerous intertidal fish and invertebrates release eggs or larvae during semilunar, large amplitude, nocturnal tides when these early life stages are best able to escape predation by fish that feed near the shore during the day. Remarkably, some species, including the fiddler crabs Uca terpsichores and Uca deichmanni, maintain this timing throughout the year as temperature, and thus the rate of embryonic development, vary. The mechanisms that allow such precision in the timing of the production of young are poorly known. A preliminary study suggested that when temperature decreases, U. terpsichores mate earlier in the tidal amplitude cycle such that larvae are released at the appropriate time. We tested this idea by studying the timing of courtship in U. terpsichores and U. deichmanni as temperature varied annually during two years, at 5 locations that differed in the temperature of the sediment where females incubate their eggs. Uca terpsichores courted earlier at locations where sediment temperature declined seasonally but not where sediment temperature remained elevated throughout the year. In contrast, clear shifts in courtship timing were not observed for U. deichmanni despite variation in sediment temperature. We discuss other mechanisms by which this species may maintain reproductive timing. These two species are likely to be affected differently by changes in the frequency and intensity of cold periods that are expected to accompany climate change.     

RXR isoforms and endogenous retinoids in the fiddler crab, Uca pugilator

The pleiotropic effects of circulating ecdysteroids in the adult fiddler crab, Uca pugilator, during molting, regeneration, and reproduction are mediated by a limited number of receptor proteins. We hypothesize that hormonal effects in vivo may be the result of complex interactions between at least two receptor heterodimer conformations that differentially respond to multiple ecdysteroid/retinoid signals. Two splicing variants of the fiddler crab retinoid-X-receptor (UpRXR) differ from one another by the addition of a 33 amino acid insert in the ligand-binding domain. We show here that the ecdysteroid receptor in the fiddler crab (UpEcR) behaves differently depending upon the UpRXR isoform with which it is partnered. The two UpRXR variant partners for UpEcR confer slightly different responses in the binding of Ponasterone A (PA)—a naturally occurring ecdysteroid in the blood of Uca. UpRXR can bind 9-cis retinoic acid (9cRA) as well as terpenoids. 9cRA and the naturally occurring terpenoid, methyl farnesoate, influence the binding of PA to UpEcR/UpRXR dimers. Endogenous retinoids are found in the blastema of regenerating limbs of Uca and they (plus blood-borne terpenoids) may add additional levels of differential response by target tissues. Thus, the two sets of heterodimers tested here may represent different dynamic complexes whose properties are defined by the specific heterodimeric subunits involved and the specific ligands available.     

Neurosecretion in the sinus gland of the fiddler crab,Uca pugnax

The ultrastructure of the sinus gland of the fiddler crab, Uca pugnax, was investigated and found to be similar to that in other crustaceans. Five types of neurosecretory axon terminals were tentatively identified on the basis of the size, shape, and electron density of granules within the axons. Release of neuro-secretory material appears to be by exocytosis.     

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.     

Female Choice in the Synchronously Waving Fiddler Crab Uca annulipes

In the fiddler crab, Uca annulipes, males attract receptive females into their burrows by waving their greatly enlarged major claw. We have previously shown that males clustered around a female wave in close synchrony. Females may have a preference for leading signals and synchronised waving may arise as an epiphenomenon of competition between males to signal first. Indeed, the males in clusters that females approach and visit in their burrows are more likely to produce leading waves than are their neighbours. Here we document two other differences in the waving behaviour of visited males and their neighbours. First, visited males complete the downward component of the wave more rapidly than their neighbours. Second, the interval between the end of one wave and the start of the next is shorter for visited males. How can waving be synchronous if visited males wave faster than their neighbours? While only 9% (40/431) of waves by neighbours did not overlap those of the visited male, 22% (110/501) of visited male waves did not overlap the wave of a focal neighbour (111 visited male-neighbour dyads). Hence, while overlapping waves are nearly synchronous, visited males produce additional, ‘nonoverlapping’ waves that result in a higher wave rate than that of their neighbours.     

A preference for a sexual signal keeps females safe

Predation is generally thought to constrain sexual selection by female choice and limit the evolution of conspicuous sexual signals. Under high predation risk, females usually become less choosy, because they reduce their exposure to their predators by reducing the extent of their mate searching. However, predation need not weaken sexual selection if, under high predation risk, females exhibit stronger preferences for males that use conspicuous signals that help females avoid their predators. We tested this prediction in the fiddler crab Uca terpsichores by increasing females' perceived predation risk from crab-eating birds and measuring the attractiveness of a courtship signal that females use to find mates. The sexual signal is an arching mound of sand that males build at the openings of their burrows to which they attract females for mating. We found that the greater the risk, the more attractive were males with those structures. The benefits of mate preferences for sexual signals are usually thought to be linked to males' reproductive contributions to females or their young. Our study provides the first evidence that a female preference for a sexual signal can yield direct survival benefits by keeping females safe as they search for mates.     

Morphology and function of the reproductive system of representatives of the genus Uca

The morphology of the reproductive organs of three species of fiddler crabs, Uca ecuadoriensis, Uca c.f. forcipata, and Uca tangeri were investigated to subsequently produce a model of their mode of operation. Vulva, vagina, and spermatheca in females, and the first and second gonopods in males were examined by applying histological techniques and electron microscopy. In all three species, vulva and vagina conform to the concave type, and the spermatheca complies with the ventral type. The tissue of the oviduct orifice is enlarged and bulges into the lumen of the spermatheca. Differences between the three species are apparent in the organization of the spermatheca, especially in the distribution and structure of glandular epithelium: In U. ecuadoriensis and U. c.f. forcipata the largest proportion of the spermathecal wall is lined with cuticle and only a small area consists of glandular epithelium, while in U. tangeri almost all of the lining is glandular. Furthermore, the glandular epithelia of the species differ in their histology and ultrastructure: In U. ecuadoriensis it is tubular and multilayered, while in U. c.f. forcipata it is mono‐layered. U. tangeri finally has both forms of this tissue. In the males, the terminal segments of the first gonopod exhibit a tight fit to female organs and narrow, tightly sealed sperm channels. These features suggest a tendency towards minimizing loss of fluids, which can be interpreted as an adaptation to mating on land. The tight fit of male gonopod and female opening seem to be protection from interbreeding, which points toward a strong sexual selection. In the terrestrial environment, these originally aquatic organisms experience serious competition for resources; therefore there is pressure on successful reproduction. According to the current results a model of the process of fertilization and egg‐laying involving the investigated organs was generated. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.