Pillar Function in the Fiddler Crab Uca beebei (II): Competitive Courtship Signaling

Male Uca beebei court and attract females into burrows they defend on muddy sand flats in the intertidal zone on the Pacific coast of the tropical Americas. Mating, oviposition and incubation (breeding) occur underground in males' burrows. Some courting males build mud pillars (2 cm high) at the entrance of their burrow. The purpose of this field study was to assess the role of pillars in competitive courtship signaling among males. I studied the effect of pillars on female behavior by recording the responses of wandering females to courtship from males resident at burrows with and without pillars. I also caught females, released them individually in a circular arena with an equal number of empty burrows with and without pillars around its circumference, and chased the females with a simulated avian predator. Females moved to burrows of both types more often when they were courted (82%) than when they were chased (67%). Receptive females were attracted to the burrows of the males that courted them significantly more often (97%) when these burrows had pillars than when they lacked pillars (66%). However, once females entered males' burrows they were equally likely to remain, mate and breed in both types of burrows. Females also more often moved to burrows with pillars (66%) than to burrows without pillars when they ran from the simulated predator. Both male courtship displays and pillars probably provide cues females use to locate males' burrows. The visual similarity between pillars and a display courting males give immediately before they enter their burrows suggests that pillars are icons of the display. The effect of pillars on female behavior, the timing of pillar building relative to when females choose mates, and contrasts in the behavior of males that do and those that do not build pillars suggest that pillar building has evolved due to competition among males to attract females into their burrows.     

Locomotor Rhythms in the Fiddler Crab, Uca subcylindrica

The locomotor activities of individual specimens of Uca subcylindrica (Stimpson) collected from semi-arid, supratidal habitats in south Texas and northeastern Mexico were studied in the laboratory using periodogram analysis. When crabs were placed under constant darkness (DD) or constant illumination (LL), free-running circadian rhythms were observed in the activity recordings. The locomotor activity of strongly rhythmic crabs in LL has an average period length of 24.4 h. Crabs held in DD express motor rhythms with periods of approximately 24.0 h. In LL the most common wave form for activity is unimodal, while under DD it is bimodal. Recordings under natural illumination (NL) revealed that both period length and the time of maximum activity (phasing) varied through the year. During winter months, the crabs are primarily diurnal with peaks in activity occurring between 0900 and 2100 h and possess a circadian rhythm with a 23.9 h period. During summer, crabs were nocturnal with maximal activity between 1300 and 0600 and a circadian period closer to 24.0 h. In these experiments, the rhythmic locomotor activities of U. subcylindrica are best described as “circadian”. This is unusual for a genus known for its expression of circatidal and circalunidian rhythms.     

Locomotor Rhythms in the Fiddler Crab,Uca subcylindrica

The locomotor activities of individual specimens of Uca subcylindrica (Stimpson) collected from semi-arid, supratidal habitats in south Texas and northeastern Mexico were studied in the laboratory using periodogram analysis. When crabs were placed under constant darkness (DD) or constant illumination (LL), free-running circadian rhythms were observed in the activity recordings. The locomotor activity of strongly rhythmic crabs in LL has an average period length of 24.4 h. Crabs held in DD express motor rhythms with periods of approximately 24.0 h. In LL the most common wave form for activity is unimodal, while under DD it is bimodal. Recordings under natural illumination (NL) revealed that both period length and the time of maximum activity (phasing) varied through the year. During winter months, the crabs are primarily diurnal with peaks in activity occurring between 0900 and 2100 h and possess a circadian rhythm with a 23.9 h period. During summer, crabs were nocturnal with maximal activity between 1300 and 0600 and a circadian period closer to 24.0 h. In these experiments, the rhythmic locomotor activities of U. subcylindrica are best described as “circadian”. This is unusual for a genus known for its expression of circatidal and circalunidian rhythms.     

Regeneration of Walking Legs in the Fiddler Crab Uca pugilator

SYNOPSIS. Regeneration of walking legs in the fiddler crab Ucapugilator is most efficient when it follows autotomy (the reflexiveloss of a limb). Closure of the wound and would healing occurimmediately following autotomy and visible regeneration beginswithin a few days. Regeneration of the walking leg occurs intwo distinct stages: The first stage, called Basal Growth, involvesmitosis and differentiation. The second stage involves primarilyprotein synthesis and water uptake and is called ProecdysialGrowth. Proecdysial Growth is, in part, under direct hormonalstimulation.     

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.     

Hormonal Control of the Molt Cycle in the Fiddler Crab Uca pugilator

Comparison of the circulating steroids in the blood of crabslacking neurosecretory eyestalk centers and crabs with thosecenters intact (but lacking several walking legs) has revealeddifferences in vivo that can be attributed to eyestalk factors.It is concluded that eyestalk factors (including the putativemolt-inhibiting hormone, MIH): 1) exert control in vivo overproduction of 25-deoxyecdysone by the crab Y-organ, 2) controlcyclic steroid production throughout anecdysis and parts ofproecdysis and 3) are not solely responsible for the increasedsteroid production that occurs during late proecdysis. Usingthe data presented here and previously published data, a simplefeedback model for the control of Y-organ activity is proposed.The model suggests that the Y-organs of the crabUca pugilatorare modestly active during anecdysis but become further activatedduring late proecdysis. The increased activation requires morethan eyestalk removal and may involve additional extra-eyestalkfactors.     

Limb Regeneration in the Fiddler Crab, Uca pugilator: Hormonal and Growth Factor Control

This paper summarizes recent work on various aspects of hormonalcontrol of regeneration in the crustacean, Uca pugilator. Hormonalcontrol in this crab is effected by means of the crustacteansteroid hormones, the ecdysteroids. New evidence is presentedsupporting a role for the retinoid hormones, all-trans retinoicacid and 9-cis retinoic acid, in the control of regenerationin these animals. The possible role of fibroblast growth factorsin organization of the limb blastema is explored and the similaritiesbetween vertebrate and invertebrate control of regenerationare discussed.     

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.     

Limb Regeneration in the Fiddler Crab, Uca pugilator: Histological, Physiological and Molecular Considerations

This paper summarizes our recent work on the histological, physiologicaland molecular aspects of limb regeneration in the fiddler crabUca pugilator and new information is presented on mitotic activityin the blastema during the first days of blastemal organization.We also report for the first time the localization of vertebrategrowth factor immunoreactivity (FGF 2 and FGF 4) in the regeneratingblastema. In the first part of this paper we review recent histologicalfindings concerning the physical events that accompany autotomyof limbs and propose a new function for the autotomy membrane—thetethering of the regenerating pedal nerve to the walls of thecoxa. In the second part of the paper we review our recent findingson the identification and characterization of the Uca ecdysteroidreceptor (UpEcR, and its potential dimer partner, the retinoid-X-receptor,UpRXR). Using Uca-specific antibody probes raised in our lab,we have been able to identify specific cells in the early blastemathat express receptor proteins. The regenerating limb of thefiddler crab is responsive to both steroids and retinoids andmRNA for steroid and retinoid receptors are expressed in theregenerating limb buds during all stages of regeneration. TheDNA and deduced amino acid sequences of the ecdysteroid receptoris very similar to the sequences of insect EcRs, while the retinoidreceptor is similar to insect protein (ultraspiracle) in theDNA-binding domain, but closer to vertebrate RXRs in the ligandbinding domain     

Locomotor Activity of the Fiddler Crab,Uca subcylindrica (Stimpson), under Artificial Illumination

Specimens of the fiddler crab Uca subcylindrica (Stimpson) were captured in south Texas (USA) for locomotor rhythm studies. Actographic data were analyzed using Tau? sofware. Under constant illumination (LL) and darkness (DD), the semiterrestrial crabs express a circadian rhythm of locomotion. When exposed to illumination/darkness cycles (LD_12:12 or LD_14:10), their bouts of activity are entrained to the photoperiod. In LD, activity is generally bimodal with an initial burst about 0.5 h after illumination. A second burst of activity begins 1 to 2 h before the end of illumination. When transferred from LD to LL, a locomotor rhythm with an average period of 24.6 ± 1.0 h (n = 19) is expressed in 89 percent of the crabs. On the other hand, when placed in DD after LD (n = 8), the crabs are either arrythmic or weakly rhythmic (period = 23.8 ± 0.2 h; n = 2). If the onset of illumination is advanced by 6 h, a period of less than 24.0 h is detected in the actogram. If the onset of illumination is delayed by 6 h, a locomotor rhythm with a period greater than 24.0 h appears. The locomotor behavior of this species of fiddler crab, Uca subcylindrica, is not related to the tidal rhythmicities seen in other members of the genus Uca. Rather, it has strong circadian components.     

Locomotor Activity of the Fiddler Crab, Uca subcylindrica (Stimpson), under Artificial Illumination

Specimens of the fiddler crab Uca subcylindrica (Stimpson) were captured in south Texas (USA) for locomotor rhythm studies. Actographic data were analyzed using Tau™ sofware. Under constant illumination (LL) and darkness (DD), the semiterrestrial crabs express a circadian rhythm of locomotion. When exposed to illumination/darkness cycles (LD_12:12 or LD_14:10), their bouts of activity are entrained to the photoperiod. In LD, activity is generally bimodal with an initial burst about 0.5 h after illumination. A second burst of activity begins 1 to 2 h before the end of illumination. When transferred from LD to LL, a locomotor rhythm with an average period of 24.6 ± 1.0 h (n = 19) is expressed in 89 percent of the crabs. On the other hand, when placed in DD after LD (n = 8), the crabs are either arrythmic or weakly rhythmic (period = 23.8 ± 0.2 h; n = 2). If the onset of illumination is advanced by 6 h, a period of less than 24.0 h is detected in the actogram. If the onset of illumination is delayed by 6 h, a locomotor rhythm with a period greater than 24.0 h appears. The locomotor behavior of this species of fiddler crab, Uca subcylindrica, is not related to the tidal rhythmicities seen in other members of the genus Uca. Rather, it has strong circadian components.     

氨氮对不同生长阶段的弧边招潮急性毒性研究

在pH8.09、水温28℃±1℃、盐度5‰条件下,进行了氨氮对不同生长阶段的弧边招潮急性毒性试验。结果显示:氨氮对全甲宽小于1.5cm、1.5cm、1.5cm至2.5cm之间3个生长阶段的弧边招潮96h的半数致死浓度(LC50)分别为47.28mg·L-1、71.49mg·L-1和90.57mg·L-1,三者之间差异极显著(P<0.01),对应的非离子氨浓度为3.71mg·L-1、5.61mg·L-1和7.11mg·L-1;氨氮的安全浓度分别为4.73mg·L-1、7.15mg·L-1和9.06mg·L-1,对应的非离子氨浓度为0.37mg·L-1、0.56mg·L-1和0.71mg·L-1。结果表明,氨氮对弧边招潮的毒性与氨氮浓度和中毒时间呈正相关。3个生长阶段弧边招潮对氨氮的耐受性大小次序为:全甲宽1.5cm至2.5cm之间>全甲宽1.5cm>全甲宽小于1.5cm。     

Variation in the Boldness of Courting Sand Fiddler Crabs (Uca pugilator)

Individuals can express boldness in their readiness to resume courtship signaling following a perceived threat. The degree of boldness that is selectively favored depends on the magnitude of costs and benefits that may vary across time and space. We examined within‐ and between‐individual variation in the boldness of courting male sand fiddler crabs, Uca pugilator, across an entire breeding season at a South Carolina (USA) salt marsh where courtship is restricted to supratidal embankments. Boldness was assessed by the time to re‐emergence and the number of re‐emergences of males who were purposely startled into their breeding burrows once every 3 min for a total of five times. The two measures of boldness were significantly positively correlated. Courting males are on average bolder when their density is high and when tidal conditions correspond to peaks in the number of females moving over the embankment surface. Time to re‐emergence increases with successive startles although some males consistently re‐emerge faster than others. Large males are not bolder than small males. When male density is high, nearest neighbors frequently re‐emerge at the same time, suggesting that males cue on the responses of other nearby males, perhaps by assessing substrate vibration. This may reduce the chance of losing a potential mate to a local competitor.     

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.     

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.     

Contextual Cross-Referencing of Species Names for Fiddler Crabs (Genus Uca): An Experiment in Cyber-Taxonomy

Cyber-taxonomy of name usage has focused primarily on producing authoritative lists of names or cross-linking names and data across disparate databases. A feature missing from much of this work is the recording and analysis of the context in which a name was used—context which can be critical for understanding not only what name an author used, but to which currently recognized species they actually refer. An experiment on recording contextual information associated with name usage was conducted for the fiddler crabs (genus Uca). Data from approximately one quarter of all publications that mention fiddler crabs, including 95% of those published prior to 1924 and 67% of those published prior to 1976, have currently been recorded in a database. Approaches and difficulties in recording and analyzing the context of name use are discussed. These results are not meant to be a full solution, rather to highlight problems which have not been previously investigated and may act as a springboard for broader approaches and discussion. Some data on the accessibility of the literature, including in particular electronic forms of publication, are also presented. The resulting data has been integrated for general browsing into the website http://www.fiddlercrab.info; the raw data and code used to construct the website is available at https://github.com/msrosenberg/fiddlercrab.info.     

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.     

Reproductive Aspects of a Tropical Population of the Fiddler Crab Uca annulipes (H. Milne Edwards, 1837) (Brachyura: Ocypodidae) at Costa Do Sol Mangrove, Maputo Bay, southern Mozambique

Uca annulipes(H. Milne Edwards) is the dominant fiddler crab inhabiting mangrove forests in East Africa. However, several aspects concerning its reproduction are poorly known. Such information is necessary for a better understanding of several biological aspects occurring in this species. The aim of this study was to assess the reproductive biology of Uca annulipes as well as some factors determining its breeding season. Samplings were undertaken from January to December 2002 at Costa do Sol, a tropical mangrove forest around Maputo Bay, southern Mozambique. Crabs were randomly sampled, measured for carapace width (CW) and checked for the presence of eggs on female pleopods. Like most tropical brachyurans, Uca annulipes follows a continuous reproduction throughout the year with peaks of spawning during summer season and low frequency of egg-bearing females during winter. The frequency of ovigerous females was correlated with the air temperature (r ²= 0.652, p < 0.05) and rainfall (r ²= 0.897, p < 0.001), with the last parameter being the most important factor regulating the reproductive activity of this crab.     

Determinants of Male Spacing Behaviour in Panacanthus pallicornis (Orthoptera: Tettigoniidae)

Panacanthus pallicornis is a neotropical species of katydid endemic to Colombia that inhabits premontane forests, and individuals are found in regenerating (RF) and dense forest (DF) habits. Members of this species are made conspicuous by their colouration, remarkable defensive body thorns and the loud mating calls of the males. Through capture, marking, and recapture, we determined singing‐site characteristics in two different natural habitats: an RF and a DF. Using nearest neighbour analysis, we found male distribution tends to uniformity in forest habitats, but in the RF, male distribution is more random. Males also showed preference for high singing sites and these were correlated with host plant height. Although host plants in DF were taller than those in the RF, male’s perch preference was independent from the habitat; and in RF, males select lower perches. To investigate if male spacing was a function of the calling song, we manipulated two groups of males from a different population, one group deafened (tympanic membranes torn), and another control group with tympanic membranes intact. Insects were released from a single location in the two different zones of similar area. After a number of days we measured male displacement, and found that the deafened group distribution tended to aggregation, while the control group spread throughout the experimental zone with a random distribution. These results suggest that male spacing behaviour is one of the functions of the calling song. Based on a pre‐established phylogenetic framework of the genus Panacanthus, we discuss some implications for the evolution of the calling song of P. pallicornis in both RF and DF.     

Aggression among Lekking Male Fallow Deer (Dama dama): Territory Effects and Relationship with Copulatory Success

We studied the aggressive behavior of territorial male fallow deer (Dama dama) at two leks. Daily number of escalated fights was not correlated with number of matings. A dominance index including all agonistic interactions was weakly correlated with copulatory success; the correlation was stronger when each half of the rut was considered separately. Dominance likely changed over the rut due to fatigue. We ranked lek territories by the number of copulations seen in each. Males that won fights were likely to take over the loser's territory only if it ranked higher than the one they held. Winners were more likely to fight with a third male after defeating an opponent that held a higher-ranking territory than one from a lower-ranking territory. Exposure to attacks by third-party males may be a major cost of fighting. Males appear aware of the relative value of different lek territories, but the frequency of aggression was not scaled to potential fitness benefits, possibly because males seldom fought with opponents they were unlikely to beat. The outcome of interactions on the lek may not always reflect the relative dominance rank of the contestants. Females are unlikely to use the outcome of fights as a direct criterion for mate selection.