Territorial battles between fiddler crab species.

Many species worldwide are impacted by habitat loss. This may result in increased competition both within species and between species. Many studies have demonstrated that when two previously non-overlapping species are forced to compete over a resource, one species is likely to become dominant over the other. This study explores the impact a larger species of fiddler crab (Tabuca elegans-previously known as Uca elegans) has when invading an area previously used solely by a smaller species (Austruca mjoebergi-previously known as Uca mjoebergi). Here we show that, while there are some detrimental effects of living next to a heterospecific, they are relatively minor. New heterospecific neighbours fight more regularly with resident crabs, but each fight is no longer or more escalated than those between the resident and a new conspecific male. The residents are not specifically targeted by intruding heterospecifics, thus, given the large advantage of having a heterospecific neighbour in terms of lowered competition for females, the overall impact of species mixing is probably not as negative as might have been predicted.

Not what it looks like: mate-searching behaviour, mate preferences and clutch production in wandering and territory-holding female fiddler crabs.

Risks inherent in mate-searching have led to the assumption that females moving sequentially through populations of courting males are sexually receptive, but this may not be true. We examined two types of fiddler crab females: wanderers moving through the population of courting males and residents that were occupying and defending their own territories. Sometimes residents leave territories to look for new burrows and we simulated this by displacing wanderers and residents and observing their behaviour while wandering. We predicted that the displaced wanderers would exhibit more mate-searching behaviours than resident females. However, wandering and resident females behaved nearly identically, displaying mate-searching behaviours and demonstrating matching mate preferences. Also, males behaved the same way towards both female types and similar proportions of wanderers and residents stayed in a male's burrow to mate. But more wanderers than residents produced egg clutches when choosing a burrow containing a male, suggesting females should be categorized as receptive and non-receptive. Visiting and rejecting several males is not the defining feature of female mate choice. Moving across the mudflat by approaching and leaving a succession of burrows (mostly occupied by males) is an adaptive anti-predator behaviour that is useful in the contexts of mate-searching and territory-searching.

Mate choice and the operational sex ratio: an experimental test with robotic crabs.

The operational sex ratio (OSR: sexually active males: receptive females) predicts the intensity of competition for mates. It is less clear, however, under what circumstances, the OSR predicts the strength of sexual selection - that is, the extent to which variation in mating success is attributable to traits that increase the bearer's attractiveness and/or fighting ability. To establish causality, experiments that manipulate the OSR are required. Furthermore, if it is possible to control for any OSR-dependent changes in the chosen sex (e.g. changes in male courtship), we can directly test whether the OSR affects the behaviour of the choosing sex (e.g. female choice decisions). We conducted female mate choice experiments in the field using robotic models of male fiddler crabs (Uca mjoebergi). We used a novel design with two females tested sequentially per trial. As in nature, the choice of the first female to mate therefore affected the mates available to the next female. In general, we detected significant sexual selection due to female choice for 'males' with larger claws. Importantly, the strength of sexual selection did not vary across five different OSR/density treatments. However, as the OSR decreased (hence the number of available males declined), females chose the 'males' with the largest claws available significantly more often than expected by chance. Possible reasons for this mismatch between the expected and observed effects of the OSR on the strength of sexual selection are discussed.

Why do fiddler crabs build chimneys?

Chimneys are mud mounds built by fiddler crabs that encircle the entrance to their burrow. Their function in many species is unknown. In Uca capricornis, crabs of both sexes and all sizes build chimneys, but females do so disproportionately more often. There are no differences in the immediate physical or social environments between crabs with and without a chimney. Chimney owners spend less time feeding and more time underground than non-owners. We show experimentally that burrows with a chimney are less likely to be located by an intruder. It is possible that some crabs construct chimneys around their burrow to conceal the entrance and reduce the risk of losing it to an intruder.

What factors contribute to an ownership advantage?

In most taxa, owners win fights when defending a territory against intruders. We calculated effect sizes for four factors that potentially contribute to an 'owner advantage'. We studied male fiddler crabs Uca mjoebergi, where owners won 92% of natural fights. Owners were not more successful because they were inherently better fighters (r=0.02). There was a small effect (r=0.18) of the owner's knowledge of territory quality (food availability) and a medium effect (r=0.29) of his having established relations with neighbours (duration of active tenure), but neither was statistically significant. There was, however, a significant effect due to the mechanical advantage the owner gained through access to the burrow during fights (r=0.48, p<0.005).

What are the consequences of being left-clawed in a predominantly right-clawed fiddler crab?

Male fiddler crabs (genus Uca) have an enlarged major claw that is used during fights. In most species, 50% of males have a major claw on the left and 50% on the right. In Uca vocans vomeris, however, less than 1.4% of males are left-clawed. Fights between opponents with claws on the same or opposite side result in different physical alignment of claws, which affects fighting tactics. Left-clawed males mainly fight opposite-clawed opponents, so we predicted that they would be better fighters due to their relatively greater experience in fighting opposite-clawed opponents. We found, however, that (i) a left-clawed male retains a burrow for a significantly shorter period than a size-matched right-clawed male, (ii) when experimentally displaced from their burrow, there is no difference in the tactics used by left- and right-clawed males to obtain a new burrow; however, right-clawed males are significantly more likely to initiate fights with resident males, and (iii) right-clawed residents engage in significantly more fights than left-clawed residents. It appears that left-clawed males are actually less likely to fight, and when they do fight they are less likely to win, than right-clawed males. The low-level persistence of left-clawed males is therefore unlikely to involve a frequency-dependent advantage associated with fighting experience.

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.