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.     

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.     

Causes of a male-biased operational sex ratio in the fiddler crab<Emphasis Type="Italic"> Uca crenulata</Emphasis>

This research investigates the causes of a male-biased operational sex ratio in a population of the California fiddler crab, Uca crenulata. Mensurative studies revealed there were almost twice as many adult males as females, mating occurred across half of the days within the breeding season, and females had much longer individual reproductive cycles than males. Therefore, many more males than females were available for mating on each breeding day. Perhaps as a consequence, males spent a large proportion of their time fighting with neighbors and rapidly waving their large claws when females passed by. Electronic Publication     

The Function of the Four Types of Waving Display in Uca lactea: Effects of Audience,Sand Structure,and Body Size

Multiple signals that convey different messages have been reported in many taxa, but relatively few studies have been made on such signals in invertebrates. In the present study, I investigated four types of claw‐waving display used in the fiddler crab Uca lactea to test whether the displays have different functions. Three males with a sand structure beside their burrows (which can attract females) and three males without a sand structure were fenced in an opaque enclosure, and I videotaped their waving displays after releasing two burrowless males or two burrowless females to test the effects of audiences. (a) Lateral‐circular waving tended to occur in enclosures with burrowless females and was performed frequently by males that had sand structures. (b) Lateral‐flick waving was performed frequently by males without sand structures, and its frequency was positively correlated with the signaler’s body size. (c) Rapid‐vertical waving was observed frequently in enclosures with burrowless males, and its frequency was negatively correlated with the signaler’s body size. (d) Circular waving tended to occur in enclosures with burrowless females and was performed frequently by males that had sand structures, and its frequency was positively correlated with the signaler’s body size. In my previous study, lateral‐circular waving was often seen in the breeding season and was mostly performed to female audiences, lateral‐flick waving was frequently performed to neighboring resident males, rapid‐vertical waving was performed mainly to intruding burrowless males, and circular waving did not have apparent audiences in most cases. Finally, I concluded that lateral‐circular waving was used as a courtship display, lateral‐flick waving was related to border disputes, rapid‐vertical waving was used for burrow guarding, and circular waving was used to broadcast the signaler’s general quality.     

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.     

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.     

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     

For whom the male waves: four types of claw-waving display and their audiences in the fiddler crab, <Emphasis Type="Italic">Uca lactea</Emphasis>

Some animals are known to use several different signals which convey different messages. In the fiddler crab, Uca lactea, I found that males performed at least four types of claw-waving display: lateral-circular, lateral-flick, rapid-vertical, and circular waving. The major audiences and the seasonal occurrence patterns of the displays differed among waving types. Lateral-circular waving (combinations of slow lateral extension, quick flexion, and circumduction of the major claw) was mostly performed to female audiences and was observed frequently in the breeding season. Lateral-flick waving (quick lateral abduction to the audience) was frequently performed to neighbor residents but rarely performed to females in the breeding season. Rapid vertical waving (rapid dorso-ventral protraction and retraction) was observed throughout the observation period and was most frequently performed to burrowless males. Circular waving (simple circumduction) was frequently observed prior to the breeding season and had no obvious audience in most cases. The results showed that males performed different types of claw-waving in different contexts. Males may have needed to use several different types of waving in order to transmit different messages. Digital video images relating to this article are available at , , , .     

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.     

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.     

The effect of claw size and wave rate on female choice in a fiddler crab

How do females select a mate when they have mating preferences for multiple male traits? In experimental studies, female fiddler crabs (Uca mjoebergi) show a strong preference for males with larger claws and higher wave rates. In the field, there is no correlation between male claw size and observed wave rate. Here we document natural mating behaviour and show that females approach males who wave at a higher rate than nearby competitors. On average, an approached male had a significantly larger claw than his two nearest neighbours but did not differ in size from his two closest waving competitors. In general, smaller males were less likely to wave at approaching females. Females therefore approached mates based directly on wave rate but, because smaller males were less likely to wave, this indirectly resulted in female choice for larger than average males. Our study raises two issues. First, how do we relate the field results to previous experimental studies showing a female preference for larger claws? Second, in U. mjoebergi, males defend smaller neighbours against intruders. Our study suggests that one benefit of such defence coalitions is to decrease the number of immediate competitors present during female mate choice by retaining smaller neighbours.     

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.     

Variation in courtship rate in the fiddler crab Uca annulipes: is it related to male attractiveness?

We investigated among-male variation in courtship waving inthe fiddler crab Uca annulipes. Wave rate is positively correlatedwith both male carapace size and relative claw size (controlledfor body size), and relative claw size is positively correlatedwith an index of body condition. An experimental reduction inthe availability of food decreased male wave rate. These datasuggest that some of the variation in wave rate among malesis due to variation in male condition combined with energeticcosts to waving (differential costs). However, we also foundthat the correlation between male size and wave rate decreasedover the semilunar cycle. Later in the cycle, smaller malesincrease their wave rate relative to that of larger males. Previouswork has shown that females are more likely to accept a smallermale as a mate later in the cycle. We suggest that smaller malesinvest disproportionately more in courtship later in the cyclebecause the potential benefits are greater due to their increasedattractiveness to females (differential benefits). Alternativeexplanations for the observed temporal trend are also discussed.     

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.     

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.     

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).     

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.     

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     

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

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

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.