The evolution of waving displays in fiddler crabs (Uca spp., Crustacea: Ocypodidae)

Male fiddler crabs are commonly recognized by the presence of a single massive claw used in a variety of contexts, including territorial defence, agonistic interactions, and courtship behaviour. The most common behavioural context involving these enlarged chelipeds is their use in waving displays, which are remarkably diverse among species. Although the waving display is one of the most obvious behavioural features of male fiddler crabs, little is known about their main evolutionary trends during the diversification of the genus. The present study employed phylogenetic comparative methods to investigate the evolution of waving behaviour in a sample of 19 species of Uca from Central and North America. Digital recordings were used to quantify the temporal dynamics of waving behaviour in each species. Multivariate ordination methods were used to assess whether different elements of the display showed distinct evolutionary dynamics, particularly with respect to body size and the environment where species are most commonly found. Most of the interspecific variation in displays involves differences in the overall waving velocity, with no correspondence to their local environments, nor their body size. Interestingly, despite the strong concentration of variance in the first two ordination axes, there was no statistically significant evidence for phylogenetic signals in their respective scores. These results suggest that the overall structure of waving displays is evolutionarily labile, at the same time as being concentrated in a few particular axes of variation, possibly indicating evolution along lines of least resistance. The approach employed in the present study highlights the utility of phylogenetic comparative methods for elucidating the evolution of complex behavioural characteristics, such as the waving display in male fiddler crabs. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 307–315.     

Are females of Ilyoplax pusilla (Brachyura: Dotillidae) attracted to groups having more waving males?

Males of the dotillid crab Ilyoplax pusilla wave at approaching females during the breeding season. They also, however, often perform waving that is not directed toward any particular individual. This undirected waving is associated with the presence of male neighbors and may function in male–male competition. It may also, however, act as a long-range female attractant. To test whether undirected waving functions to attract females, we conducted a field experiment that manipulated the abundance of waving males. We found that females preferred to approach groups that had more waving males. This suggests that undirected waving by male I. pusilla functions as a long-range courtship signal.     

Coordination and neuromuscular control of rhythmic behaviors in the blue crab,Callinectes sapidus

The stereotypical courtship display (CD) behavior of the male blue crab, Callinectes sapidus, includes an unusual component: the rhythmic waving of the swimming appendages above the carapace. This behavior occurs in a unique context but it resembles two other rhythmic behaviors performed using the swimming legs: sideways swimming and backward swimming. As a first step to understanding the mechanisms that allow the expression of apparently different rhythmic motor patterns, we have examined these behaviors using slow motion video analysis and electromyography of the basal muscles of the swimming legs in freely behaving crabs. The results show that these behaviors are distinguished by four parameters: the frequency of leg waving, the phase relationship between the legs, the presence of a stationary pause in basal muscle activity combined with rotation of the distal leg during CD, and an extended range of motion of these legs during CD and backward swimming, relative to sideways swimming. EMG analysis revealed that during sideways swimming, the sequence of muscular activity between the two legs was different. In contrast, during CD and backward swimming the sequence of activity for these legs is identical.Abbreviations CD courtship display - EMGs electromyograms - CD AMP courtship display in crabs with amputated fifth legs - CD1 crabs that voluntarily used one leg to perform courtship display waving - CD 1–3 courtship waving in cycles 1–3 - CD MID courtship waving after cycles 1–3 - M-C meral-carpal joint     

Changes in the selection differential exerted on a marine snail during the ontogeny of a predatory shore crab

Empirical estimates of selection gradients caused by predators are common, yet no one has quantified how these estimates vary with predator ontogeny. We used logistic regression to investigate how selection on gastropod shell thickness changed with predator size. Only small and medium purple shore crabs (Hemigrapsus nudus) exerted a linear selection gradient for increased shell‐thickness within a single population of the intertidal snail (Littorina subrotundata). The shape of the fitness function for shell thickness was confirmed to be linear for small and medium crabs but was humped for large male crabs, suggesting no directional selection. A second experiment using two prey species to amplify shell thickness differences established that the selection differential on adult snails decreased linearly as crab size increased. We observed differences in size distribution and sex ratios among three natural shore crab populations that may cause spatial and temporal variation in predator‐mediated selection on local snail populations.     

Courtship tactics by male Ilyoplax pusilla (Brachyura, Dotillidae)

Mating in the dotillid crab Ilyoplax pusilla occurs after the female enters the male’s burrow in the tidal flat. Males use two tactics to cause females to enter their burrows for mating: the male either directs claw waving to the female (courting-wave display), to which the females responds by following the male to his burrow, or the male runs rapidly away from, then back toward, his burrow (dash-out-back display), which startles the female into his burrow. Males more often used the courting-wave than the dash-out-back display, but mating success did not differ between the two tactics. Male use of either tactic was influenced by date, female density and male size; the courting-wave display was used by larger males, later in the breeding period, and under higher female density.     

Stereotypy and variation in the claw waving display of the fiddler crab <Emphasis Type="Italic">Uca tangeri</Emphasis>

We measured temporal and spatial components of the waving display in a Uca tangeri population to look for inter-individual differences in male waving structure that may convey information about individual identity. We found evidence that the spatial components of wave structure, especially “Maximum amplitude” are responsible for most of the between-male variation of the display. This variation could reflect differences in individuals’ condition and/or could be used by conspecifics to discriminate amongst familiar and unfamiliar individuals.     

Synchronous waving in two species of fiddler crabs

In the fiddler crabs Uca saltitanta and Uca perplexa, males attract mates by waving their enlarged claws. We show that in both species waving is closely synchronised between neighbouring males in clusters, both in the presence of mate-searching females and in their absence. Wandering females visit those males in the cluster that produce more waves at faster wave rates. In U. perplexa, they also selectively visit those males that produce the greatest number of leading waves. Synchronous waving may be the result of a precedence effect causing male competition to produce leading signals.     

Choosing a mate in a high predation environment: Female preference in the fiddler crab Uca terpsichores

The interplay between a receiver's sensory system and a sender's courtship signals is fundamental to the operation of sexual selection. Male courtship signals that match a female receiver's preexisting perceptual biases can be favored yet the message they communicate is not always clear. Do they simply beacon the male's location or also indicate his quality? We explored this question in a species of fiddler crab Uca terpsichores that courts under elevated predation risk and that mates and breeds underground in the safety of males' burrows. Sexually receptive females leave their own burrows and are thereby exposed to avian predators as they sequentially approach several courting males before they choose one. Males court by waving their single greatly enlarge claw and sometimes by building a sand hood next to their burrow entrance. Hoods are attractive because they elicit a risk‐reducing orientation behavior in females, and it has been suggested that claw waving may also serve primarily to orient the female to the male. If the wave communicates male quality, then females should discriminate mates on the basis of variation in elements of the wave, as has been shown for other fiddler crabs. Alternatively, variation in elements of the claw waving display may have little effect on the display's utility as a beacon of the location of the male and his burrow. We filmed courting males and females under natural conditions as females responded to claw waving and chose mates. Analysis of the fine‐scale courtship elements between the males that females rejected and those they chose revealed no differences. When predation risk during courtship is high, males' courtship displays may serve primarily to guide females to safe mating and breeding sites and not as indicators of male quality apart from their roles as beacons.     

Eavesdropping in crabs: an agency for lady detection

Although conspicuous courtship displays are an effective way of attracting the attention of receptive females, they could provide valuable information to rival males on the location of these females. In fiddler crabs, males that see a receptive female wave their single, greatly enlarged claw in a highly conspicuous courtship display. We test whether other males use this courtship display to alert them to the presence of receptive females that they cannot directly see. We show that male fiddler crabs (Uca mjoebergi) eavesdrop on the courtship displays of nearby males to detect mate-searching females. This allows males to begin waving before a female becomes visible. Furthermore, males appear to adjust their waving according to the information available: eavesdropping males wave 12 times faster than non-courting males but only 1.7 times slower than males in full visual contact with the female.     

Waving Display in Females of Uca polita and of other Australian Fiddler Crabs

Since 1979 it is known that, in Australian species of Uca, female waving exists in addition to usual male display. The present paper deals mainly with female waving in U. polita studied in Darwin (North Australia). A few remarks on U. dampieri, U. vomeris, U. seismella and U. hirsutimanus are added. The species mentioned are members of two species groups or subgenera, which characterizes female waving as an ancestral (plesiomorph) trait. Frame by frame analysis of film sequences (open air shots) indicate homology of movements in the two sexes of U. polita. As in males, waving of females can be combined with locomotion on radial paths starting from the burrow entrance and the display is performed in series with a corresponding number of gestures. Unlike males, waving females mostly use both their chelipeds and tend to show shorter durations with regard to many of the waving parameters chosen. However, significant differences refer only to a limited number of parameters. The biological context of female waving was gathered from films and field observations. High intensity waving is released by conspecifics approaching from far (wanderers without burrows) and from the neighbourhood. Typically, only females and small males elicit high intensity display in a resident female. Waving normally stops in presence of larger males, especially of the male living in a resident breeding unit with the female in question. In spite of this, a pure agonistic (defensive) character of female waving is unlikely. Advertising of breeding condition seems to play a role similar to that in males. The few displaying females that exist in a given colony (about 2.5% in U. polita) show signs of special sexual excitement: brightening of carapace colours and sometimes spontaneous performance of waving, i.e. display immediately after emergence from the burrow in absence of any conspecific.     

Female preference for large waving claws in the dotillid crab <Emphasis Type="Italic">Ilyoplax pusilla</Emphasis>

Male Ilyoplax pusilla perform a waving display, a simple up-down movement of the claws during the reproductive season. Large males dedicated most of their surface activity to waving displays and gained higher mating success. On the other hand, small males infrequently performed waving displays and devoted their time exclusively to foraging. To examine female preference for the size of waving males, two female-release experiments were conducted. In the experiments, we recorded female choice between small- and large-waving claw models over short (10 cm) and long (25 cm) distances. In both the short- and long-distance choice experiments, significantly more females chose the large-claw model over the small-claw model. The following characteristics may produce obvious age-dependent sexual advertisement in I. pusilla, which grows throughout a life: (1) the strong female preference for large claws; (2) the short, 2-year lifespan that includes only two reproductive seasons; and (3) the lack of alternative mating strategies (e.g., surface mating).     

The presence of neighbors affects waving display frequency by Scopimera globosa (Decapoda, Ocypodidae)

Scopimera globosa, a small ocypodid crab, rhythmically raises and lowers its body and both chelae in a waving display when it is not interacting directly with another individual. To determine whether waving is a social signal and to deduce its possible function, we manipulated the sex and density of crabs in a field enclosure and recorded the waving frequency of males. Males with abundant female neighbors waved significantly more often than when the same males were caged with abundant male neighbors. Males caged with fewer neighbors of either sex seldom waved. Thus, males waved most in the presence of females, especially at high density, and least in the presence of other males, suggesting that waving may function in female acquisition. Received: August 18, 1998 / Accepted: October 2, 1999     

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.     

The population biology and genetics of the deep-sea spider crab,Encephaloides armstrongi Wood-Mason 1891 (Decapoda: Majidae)

Numerous specimens of the majid spider crab, Encephaloides armstrongi, were sampled from six stations (populations) between 150 and 650 m depth, on the continental slope off the coast of Oman. This extended the known geographic and bathymetric range of E. armstrongi, which is now known to occur along the continental margins of the northern Indian Ocean from the western coast of Burma to the coast of Oman. This band-like distribution is contiguous to the oxygen minimum zone in this region.The biology and genetics of populations of Encephaloides armstrongi separated by depth were studied. The overall sex ratio of the E. armstrongi sampled was male-biased (p less than 0.01; 3.3 males: 1 female; S_o = 0.538). However, sex ratio varied both between populations (p less than 0.01) and between size classes of crabs. Size frequency analysis indicated that the male and female crabs consisted of at least two instars, one between 6 and 16mm carapace length and one between 16 and 29 mm carapace length, which probably represented the terminal (pubertal) moult for most individuals. Accumulation of female crabs in the terminal instar probably caused the variation of sex ratio with size classes. Some male crabs grew to a larger size (up to 38 mm carapace length), possibly as a result of maturity at later instars.Length frequency distribution was significantly different between sexes (one-way ANOVA p less than 0.001). Within sexes, length frequency distributions varied between different populations. In both male and female Encephaloides armstrongi the individuals from a single population located at 150 m depth were significantly smaller than individuals at all other stations and were considered to represent a juvenile cohort. For female crabs no other significant differences were detected in length frequency between populations from 300 m to 650 m depth. Significant differences in length frequency were detected between male crabs from populations between 300 and 650 m depth.Horizontal starch gel electrophoresis was used to detect six enzyme systems coding for eight loci for individuals sampled from each population of Encephaloides armstrongi. Genetic identity (I) values between populations of E. armstrongi (I = 0.98-1.00) were within the normal range for conspecific populations. Observed heterozygosity (H_o = 0.080-0.146) was lower than expected heterozygosity (H_e = 0.111-0.160), but in the normal range detected for eukaryotic organisms.F-statistics were used to analyse between population (F_ST) and within population (F ) genetic structure. For both male and female E. armstrongi significant genetic differentiation was detected between the population located at 150 m depth and all other populations. Analyses of F_IS and F_ST, excluding the 150 m population indicated that for female E. armstrongi there was no significant structuring within or between populations. For male E. armstrongi significant heterozygote deficiencies were detected within populations and significant genetic differentiation between populations.The most likely explanations for the observations of the present study are: the population of Encephaloides armstrongi located at 150 m depth represented a juvenile cohort that is genetically distinct from deeper populations; female E. armstrongi formed a single population between 300 m and 650 m depth in the sampling area; male E. armstrongi were from two or more genetically distinct populations which are represented by different numbers of individuals at stations between 300 m and 650 m depth. This caused the observed significant differences in morphology (size distribition) and allele frequencies of male populations. It is likely that E. armstrongi exhibits gender-biased dispersal and that the crabs collected between 300 m and 650 m depth formed spawning aggressions. This also explains the bias in sex ratio of individuals sampled in the present study.     

A Spatially Explicit Model of Synchronization in Fiddler Crab Waving Displays

Fiddler crabs (Uca spp., Decapoda: Ocypodidae) are commonly found forming large aggregations in intertidal zones, where they perform rhythmic waving displays with their greatly enlarged claws. While performing these displays, fiddler crabs often synchronize their behavior with neighboring males, forming the only known synchronized visual courtship displays involving reflected light and moving body parts. Despite being one of the most conspicuous aspects of fiddler crab behavior, little is known about the mechanisms underlying synchronization of male displays. In this study we develop a spatially explicit model of fiddler crab waving displays using coupled logistic map equations. We explored two alternative models in which males either direct their attention at random angles or preferentially toward neighbors. Our results indicate that synchronization is possible over a fairly large region of parameter space. Moreover, our model was capable of generating local synchronization neighborhoods, as commonly observed in fiddler crabs under natural conditions.     

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     

Size-dependent mating preference of the male fiddler crab Austruca perplexa

In many animals, females prefer large males to small males, which allow large males to be choosier than small males when selecting a mate. We investigated the courtship intensity of small- and large-sized male fiddler crabs (Austruca perplexa) by examining their claw-waving rates (waves/min) towards small- and large-sized females. We found that large males showed a greater preference for large females by producing more waves/min towards them, whereas small males did not show any apparent preference for either large or small females. Moreover, the waving rate of large males was positively correlated with female size, but there was no correlation between waving rate and female size in small males. These results indicate that large males in a population become choosier and show strong mate choice, which is most likely due to their greater preference among females.     

Male courtship cycles in three species of tropical Ilyoplax crabs (Decapoda,Brachyura, Ocypodidae)

The courtship behaviour and cycles of male courtship activity and colouration of Ilyoplax orientalis, I. delsmani and I. gangetica were studied in the field in Malaysia and Thailand. Each species had a distinctive chela waving or beckoning display. Depending on species, the chelipeds, carapaces, or both of waving males blanched to white in contrast to the cryptic colour of nonwaving males and females. All three of these tropical Ilyoplax exhibited semilunar cycles in male waving activity at the colony level. It was confirmed for I. orientalis that individual males cycled each semilunar period between waving and non-waving phases and exhibited different behaviour toward females during these two behavioural phases.     

Keeping up appearances: male fiddler crabs wave faster in a crowd

Courtship displays are often energetically and temporally costly as well as highly conspicuous to predators. Selection should therefore favour signalling tactics that minimize courtship costs while maintaining or increasing signal attractiveness. In fiddler crabs, males court females by waving their one greatly enlarged claw in a highly conspicuous and costly display. Here, we investigate whether courting males adjust their wave rate, and therefore the cost of courtship, to the current level of competition. We show that display rate increases as competition increases and that when competition is removed, males reduce their display rate by 30 per cent. These results suggest that male fiddler crabs actively reduce the cost of courtship by adjusting their wave rate in response to the immediate level of competition.     

Analysis of waving and sound‐production display in the ghost crab,Ocypode stimpsoni

The ghost crab Ocypode stimpsoni displays waving and sound production. Sounds are produced by thumping the sand substratum with the major cheliped, and two types of sounds can be discriminated; one with a low frequency of about 12 Hz, called rapping, and another with a higher frequency (about double), called quivering. In our observations, a sequence of waving and sound emission would sometimes terminate abruptly, or appear as independent components but the component order never changed. The most frequently observed patterns were “waving with rapping and quivering”;, “waving with quivering”; and “quivering only”; quivering sounds being involved in more than 80% displays. Quivering sometimes occurred immediately after crabs emerged from the burrow, or when they returned to the entrance after discarding an excavated sand mass. The occurrence frequency of waving and sounds, the wave amplitude, and the frequency of the sound increased when other crabs approached.