Food availability in beach and marsh habitats and the size of the fiddler crab claw,a sexually selected weapon and signal

Males of the sand fiddler crab Uca pugilator possess a greatly enlarged claw that is used as a weapon in ritualized contests for control of breeding burrows and is waved to attract females to breeding burrows. Approximately 5400 crabs were collected along the Atlantic coast of North America at 14 localities, all of which had both beach and salt marsh habitats. Five measurements were made on each claw. Principal components analysis was used to generate a single measure of claw size from the seven correlated measures and scores of the claw. Carapace width was measured to index body size. Claw size was greater in beach than marsh habitats, controlling for body size. However, body size did not differ by habitat type. Claw size was also greater in laboratory‐reared males receiving more food, suggesting that differential access to food could influence habitat‐associated differences in claw size. Chlorophyll a concentration and total organic content, reflecting, respectively, the abundance of benthic algae and other food, were greater in beach than marsh habitats. Moreover, feeding opportunities were greater in the wetter beach habitat because crabs there, but not in marsh habitat, can feed at breeding burrows. Adult fiddler crabs continue to molt and grow in both body and claw size as they age. Energetic investment in the claw relative to the body is plastic. It appears that the availability of food can affect the amount of energy invested in the claw.     

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.     

Thermoregulation as an alternate function of the sexually dimorphic fiddler crab claw

Fiddler crabs are highly sexually dimorphic. Males possess one small (minor) feeding claw and one greatly enlarged (major) claw; females possess two small claws. The major claw is used to attract mates and for burrow defense, but it is costly for the male to possess. We tested the hypothesis that the major claw also functions as a thermoregulatory structure, a function that would allow males to spend a greater amount of time at the surface, foraging and attracting potential mates. Fiddler crabs Uca panacea were exposed to a source of radiant heat and body temperatures were monitored. Four groups of crabs were tested: intact males, males with the minor claw removed, males with the major claw removed, and females. The body temperatures of males without the major claw increased more rapidly and reached higher values than did those of males with the major claw intact, but the results from these animals were similar to those of females. These results support the hypothesized thermoregulatory function of the major claw. The major claw may function as a heat sink, transferring heat away from the body and dissipating it into the air. Enhanced thermoregulatory ability provided by the major claw may partially ameliorate the energetic costs of possessing such a large claw.     

Sexual selection and the physiological consequences of habitat choice by a fiddler crab

In mid-Atlantic salt marshes, reproductively active male sand fiddler crabs, Uca pugilator, use a single greatly enlarged major claw as both a weapon to defend specialized breeding burrows from other males and an ornament to attract females for mating. During the summer breeding season, females strongly prefer to mate with males controlling burrows in open areas high on the shore. Food availability decreases while temperature and desiccation stress increase with increasing shore height, suggesting that the timing and location of fiddler crab mating activity may result in a potential trade-off between reproductive success and physiological condition for male crabs. We compared thermal preferences in laboratory choice experiments to body temperatures of models and living crabs in the field and found that from the perspective of a fiddler crab, the thermal environment of the mating area is quite harsh relative to other marsh microhabitats. High temperatures significantly constrained fiddler crab activity on the marsh surface, a disadvantage heightened by strongly reduced food availability in the breeding area. Nevertheless, when the chance of successfully acquiring a mate was high, males accepted a higher body temperature (and concomitantly higher metabolic and water loss rates) than when the chances of mating were low. Likewise, experimentally lowering costs by adding food and reducing thermal stress in situ increased fiddler crab waving display levels significantly. Our data suggest that fiddler crabs can mitigate potential life history trade-offs by tuning their behavior in response to the magnitude of both energetic and non-energetic costs and benefits.     

Hunting and predation in a fiddler crab

Fiddler crabs are known primarily to be deposit feeders. They eat detritus, bacteria, and other small particles of organic material found in the sandy or muddy substrate on which they live. They have highly specialized mouthparts used to separate edible matter from nondigestable material. Here we provide evidence of cannibalism and predation in a fiddler crab, Uca annulipes. We additionally provide the first evidence of a fiddler crab hunting shrimp and insects. This study is an exemplary reminder that, even though an animal may have evolved highly specialized feeding traits, this need not preclude it from opportunistically acting as a generalist feeder.     

Dishonest signalling in a fiddler crab

Animal communication theory predicts that low-frequency cheating should be common in generally honest signalling systems. However, perhaps because cheats are designed to go undetected, there are few examples of dishonest signals in natural populations. Here we present what we believe is the first example of a dishonest signal which is used commonly by males to attract mates and fight sexual rivals. After losing their large claw male fiddler crabs (Uca annulipes) grow a new one which has less mass, is a less effective weapon and costs less to use in signalling than an equivalent-length claw of the original form. Males with original claws do not differentially fight males with regenerated claws even though they are likely to win. Regenerated claws effectively bluff fighting ability and deter potential opponents before they fight. During mate searching, females do not discriminate against males with low-mass, regenerated claws, indicating that they are deceived as to the true costs males pay to produce sexual signals. Up to 44% of males in natural populations have regenerated claws, a level unanticipated by current signalling theory. The apparent rarity of cheating may be an artefact of the usual difficulty of detecting cheats and dishonesty may be quite common.     

The functions of multiple visual signals in a fiddler crab

In many species, it is common for animals to have multiple signals within one channel of communication. Multiple signals may, however, be inefficient if they are redundant in nature. Identifying the functional significance of these multiple signals is therefore important if we are to understand the evolution of such elaborated behaviours. We proposed to identify the roles of movement-based multiple signals in a model animal system. Male fiddler crabs wave their sexually dimorphic enlarged claw during social interactions. Some species present multiple signals, where the level of complexity of the movement changes. Males of Austruca mjoebergi can perform a double wave consisting of a high- followed by a low-elevation lifting of the claw, or a single wave consisting of the high-elevation movement alone. We first investigated structural differences between the double and single wave types, and found that single waves were lower in elevation than double waves. We then explored the adaptive meaning of the wave types by manipulating the social context in which males wave. We found that double waves were given in all contexts and in higher proportions at long distances, suggesting a function of broadcasting male location. Single waves, on the other hand, were mainly given at close range and in the presence of conspecifics, suggesting intraspecific communication. Female presence elicited the highest number and proportion of single waves, a likely result of a female preference for higher wave rates. Finally, we point out that there is an element of interaction between wave types that deserves future attention. This paper is an important contribution to expand our understanding of the adaptive meaning of multiple visual signals and help reach a unified theory of their evolution.     

Residency and size affect fight duration and outcome in the fiddler crab Uca annulipes

We performed a field experiment to investigate the effect of carapace width, major cheliped length and burrow ownership on the righting success of male fiddler crabs (Uca annulipes) . We removed males from their burrows and released them back into the colony ( n = 82). Released males tended to initiate encounters with burrow owners slightly smaller than themselves. Several general predictions of Sequential Assessment Game models of contest behaviour were supported: (1) residents won more encounters; (2) intruders were more likely to win when larger than residents. When body size (carapace width) was controlled for, intruders with relatively large claws for their body size were more likely to win contests; (3) the duration of encounters was related to the size difference between males; (4) encounters won by the larger male were of shorter duration than those won by the smaller male; (5) encounters won by the resident tended to be of shorter duration than those won by intruders ( P = 0.07); (6) on average, encounter duration was longer when the intruder was larger than the resident. However, the encounters we documented began with seemingly costly behaviour such as pushing and the inter-locking of claws and did not unambiguously escalate from initial low cost behaviours. Sequential assessment of relative fighting ability may therefore not have been occurring. Prior visual assessment of opponents' fighting ability, followed by 'all-out fights' during physical encounters may also provide a plausible explanation for our results.     

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

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

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

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

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance (''surface mating'') or leave their own burrow and sequentially enter and leave (''sample'') courting males'' burrows, before staying in one to mate underground (''burrow mating''). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.     

The assessment game in sand fiddler crab contests for breeding burrows

The single enlarged claw of male sand fiddler crabs, Uca pugilator, is used in contests for control of breeding burrows. The larger of the two contestants has the larger claw and usually wins. Males use one or more of 10 agonistic elements that vary in intensity from a no-contact extension of the claw to the flip of an opponent. We used the sequence of elements employed and the duration of unstaged, naturally occurring contests in a South Carolina salt marsh to evaluate three models of extended contests: (1) energetic war of attrition, (2) cumulative assessment and (3) sequential assessment. Contests usually began with elements of low action intensity and often proceeded to elements of high intensity. Elements of higher intensity were correlated with both contest duration and the number of contest elements. Contest duration increased as opponents became more evenly matched in size, a result consistent with both cumulative and sequential assessment models. Variation in duration increased as the relative sizes of opponents increased, also in accordance with sequential assessment. The absolute size of the smaller contestant had no effect on contest duration, in contrast to predictions based on cumulative assessment or energetic war of attrition models. Contestants that lost a fight were more likely to engage immediately in another fight without loss of contest intensity, if their previous fight had been long and intense. This result is inconsistent with contests of endurance, such as the energetic war of attrition or the cumulative assessment game, but it is consistent with the ritualized display of strength and fighting skill. Thus, sequential assessment appears to best explain ritualized fiddler crab contests. Cumulative assessment, however, may be the appropriate model for extended, nonritualized, all-out fights. Cumulative assessment may also explain the tenure of individuals on breeding grounds where multiple engagements are likely to test endurance and tolerance to damage over a period of days. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.