Visually mediated species and neighbour recognition in fiddler crabs (Uca mjoebergi and Uca capricornis)

Mating signals are often directed at numerous senses and provide information about species identity, gender, receptiveness, individual identity and mate quality. Given the diversity of colourful body patterns in invertebrates, surprisingly few studies have examined the role of these visual signals in mate recognition. Here, we demonstrate the use of claw coloration as a species recognition signal in a fiddler crab (Uca mjoebergi). Furthermore, we show that distinct carapace colour patterns in Uca capricornis enable males to discriminate between their female neighbours and unfamiliar females. This is the first empirical evidence of the social importance of colour markings in fiddler crabs and the first example of visually mediated species and neighbour recognition in invertebrates other than insects.     

Male-like mudballing behavior of some female fiddler crabs (Uca tangeri)

At each low tide, male and female Uca tangeri remove mudballs from inside their burrows and place them on the surface. Previous studies have shown clear intersexual differences in mudball arrangements. However, we noticed that some females placed their mudballs in an arrangement similar to that of males. In this study, we investigated several factors that may have been responsible for this change in female mudballing behavior. We found no significant effect of the lunar cycle, female size and reproductive state, or burrow features. We briefly discuss the avoidance of sexual coercion or parasite modification of host behavior as possible factors. Our study shows that intersexual differences in mudballing behavior are more complex than previously thought. Received: October 18, 2000 / Accepted: May 7, 2001     

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     

Constraints on geographic variation in fiddler crabs (Ocypodidae: Uca) from the western Atlantic

A key question in evolutionary biology is how intraspecific variation biases the evolution of a population and its divergence from other populations. Such constraints potentially limit the extent to which populations respond to selection, but may endure long enough to have macroevolutionary consequences. Previous studies have focused on the association between covariation patterns and divergence among isolated populations. Few have focused on geographic variation among semi‐connected populations, however, even though this may be indicative of early selective pressures that could lead to long‐term divergence and speciation. Here, we test whether covariation in the shape of the carapace of fiddler crabs (genus Uca Leach, 1814) is important for structuring geographic variation. We find that morphological divergence among populations is associated with evolvability in the direction of divergence in only a few species. The shape of the ancestral covariation matrix in these species differs from other species in having notably more variation concentrated along fewer directions (i.e. higher eccentricity). For most species, there is some evidence that covariation has constrained the range of directions into which populations have diverged but not the degree of divergence. These results suggest that even though fiddler crab populations have diverged morphologically in directions predicted by covariation, constraints on the extent to which divergence has occurred may only be manifested in species where variation patterns are eccentric enough to limit populations’ ability to respond effectively in many directions.     

Correlating context-specific boldness and physiological condition of female sand fiddler crabs (Uca pugilator)

Consistent individual behavioral differences across ecological contexts are a recognized feature of animal populations. These differences can be expressed in two ways: context-specifically or context-generally. The former is characterized by consistent responses in one context (i.e. repeatability), whereas the latter by consistency that spans contexts (i.e. behavioral syndromes). The proximate causes of behavioral consistency remain unclear, yet there is evidence that physiology may couple the expression of some behavioral traits in unrelated contexts. We therefore explored the correlation between bold behavior of female sand fiddler crabs (Uca pugilator) and the condition of the hepatopancreas, an organ vital to crustacean metabolism and reproduction. We did this by taking replicate measurements of two risk-taking behaviors per individual in the contexts of predator avoidance and environment exploration, and examining correlations within and between these observations. We then determined the relationship of behavior with hepatopancreas mass and lipid content. Individual crabs responded consistently within each context. However, across-context correlations were absent, indicating that boldness is isolated, at least in the selected scenarios. Additionally, anti-predator and exploratory behaviors were significantly influenced by size but not linked to hepatopancreas physiology. Our results show that context-specific trait expression may occur in the absence of a physiological correlate.     

Ecophenotypic physiology: osmoregulation by fiddler crabs (Uca spp.) from the northern Caribbean in relation to ecological distribution

Nine species of Uca were collected across the northern Caribbean from 50 locations ranging in habitat osmolality from 18 to 2760?mmol?kg^?1 (mosmol). The osmoregulating ability of each species was assessed in solutions ranging from 30 to 3550?mosmol. Survivorship, lower and upper median lethal concentration and haemolymph isosmotic concentration ([ISO]) were estimated for each. A lower lethal concentration could not be calculated for some due to survival in low osmolality. By subgenus, average [ISO] for four species from the subgenus Minuca ranged from 587 to 768?mosmol and three species from the subgenus Leptuca from 805 to 881?mosmol. For species from the subgenus Boboruca and the subgenus Uca the [ISO] was 805 and 930?mosmol, respectively. These values reflect the habitat preference of each species in a subgenus. This study broadens our understanding of ecological physiology in Caribbean Uca and demonstrates intra- and inter-specific differences among tropical fiddler crabs.     

Behavioural adaptations of the fiddler crabs Uca vocans borealis (crane) and Uca lactea lactea (De Haan) for coexistence on an intertidal shore

The distribution and coexistence of the fiddler crabs U. vocans borealis and U. lactea lactea was investigated in the upper shore of a sandy beach with respect to particle size, water content and total organic carbon content of the sediments. A clear segregation of habitats between the two species was apparent. U. lactea lactea were only found in sediments with significantly lower total organic carbon content. U. lactea lactea had a higher ratio for the size of third maxilliped to body size than U. vocans borealis. There was no statistical difference in the median sediment particle size of the habitats where both species were found on the study shore. U. lactea lactea was larger on shores where U. vocans borealis were absent. Smaller U. vocans borealis individuals occupied sediments with higher water content and finer particles. Smaller individuals of U. vocans borealis also possessed fewer spoon-tipped setae on second maxilliped than their larger conspecifics, which were found mostly on coarser sediments. U. lactea lactea spent significantly more time on the surface than in the burrow during low tide when compared with U. vocans borealis. Both Uca species exhibited similar desiccation resistance. U. lactea lactea tends to keep its frontal region in close contact with sediments to maintain moisture when being exposed to air.     

Behavioural adaptations of the fiddler crabs Uca vocans borealis (crane) and Uca lactea lactea (De Haan) for coexistence on an intertidal shore

The distribution and coexistence of the fiddler crabs U. vocans borealis and U. lactea lactea was investigated in the upper shore of a sandy beach with respect to particle size, water content and total organic carbon content of the sediments. A clear segregation of habitats between the two species was apparent. U. lactea lactea were only found in sediments with significantly lower total organic carbon content. U. lactea lactea had a higher ratio for the size of third maxilliped to body size than U. vocans borealis. There was no statistical difference in the median sediment particle size of the habitats where both species were found on the study shore. U. lactea lactea was larger on shores where U. vocans borealis were absent. Smaller U. vocans borealis individuals occupied sediments with higher water content and finer particles. Smaller individuals of U. vocans borealis also possessed fewer spoon-tipped setae on second maxilliped than their larger conspecifics, which were found mostly on coarser sediments. U. lactea lactea spent significantly more time on the surface than in the burrow during low tide when compared with U. vocans borealis. Both Uca species exhibited similar desiccation resistance. U. lactea lactea tends to keep its frontal region in close contact with sediments to maintain moisture when being exposed to air.     

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.     

Adaptations for feeding on rock surfaces and sandy sediment by the fiddler crabs (Brachyura: Ocypodidae) Uca tetragonon(Herbst, 1790) and Uca vocans(Linnaeus, 1758)

Two species of fiddler crab, Uca tetragonon(Herbst, 1790) and Uca vocans(Linnaeus, 1758), which belong to the subgenus Gelasimus, dwell on rocky shores and muddy–sandy tidal flats, respectively, in Phuket Is., Thailand. We investigated their feeding ecology in relation to the morphology of their feeding organs: minor food-handling chelipeds and maxillipeds. U. tetragononfed chiefly on rocks covered by filamentous green algae. U. vocansfed on the emerged sand and in shallow water along the shoreline and in pools. While feeding, both crabs made sand pellets beneath their mouthparts and discarded them, indicating that they divided the matter scooped up with their minor chelipeds into edible and inedible fractions by using the maxillipeds in the water passing through their buccal cavity. The morphology of maxillipeds hardly differed between the two species, which means that both species are flotation-feeders. The morphology of their minor chelipeds, however, differed: the tips of the dactyl and pollex were flat in U. tetragononand pointed in U. vocans.When the minor cheliped was closed, U. tetragononhad a hemispherical space in the distal one-fourth of the gape, which was closed by the framing keratin layers and a few setae of the dactyl and pollex. On the other hand, U. vocanshad an ellipsoidal space in the distal half of the gape. We consider these morphological characters to be adaptations to the different feeding substrates for retaining more food-laden sediment. We discuss the role of the setae on the minor chelipeds on the basis of the morphological differences between populations of U. tetragononin Phuket Is. and East Africa where the crab inhabits muddy–sandy tidal flats.     

Determination of temperature preference and the role of the enlarged cheliped in thermoregulation in male sand fiddler crabs,Uca pugilator

• 1.Male Uca pugilator whose major cheliped was immersed in 3 °C water bath experienced a significant drop in T_b. Thus, the enlarged claw of male Uca pugilator may have an unexplored function: thermoregulation.
• 2.Crabs prefer warmer substrates (19–24 and 28–30 °C) over cooler (15–17 °C).
• 3.Mean selected temperature (MST) may not be an accurate reflection of T_b. Crabs in a thermal chamber preferred temperatures between 25 and 30 °C but their average T_b was 23.2 °C.

     

Developing methods for quantifying the apparent abundance of fiddler crabs (Ocypodidae: Uca) in mangrove habitats

Counting the number of individuals emerging from burrows is the most practical method for estimating the apparent abundance of Australian Uca species living in mangrove habitats. Experiments were conducted to investigate the effect on counts of quadrat design, distance of observer, quadrat size, recovery time and observational technique. Significant differences in the apparent abundance of one species were found when the subjects were within 2 m of the observer, and when a conspicuous quadrat was used. The largest quadrat tested provided the least variability in counts but an intermediate size (0.56 m²) was more practical. Most Uca active within a 30-min period emerged during the first 10 min regardless of site, species, sex or season. There was a linear correlation between scanning and continuous observation indicating that the former method could be useful when sampling time was limited. Temporal changes in the apparent abundance of Uca suggest that long-term sampling and more detailed studies will be worthwhile.     

Phylogenetic autocorrelation and heritability of geographic range size,shape and position of fiddler crabs,genus Uca (Crustacea,Decapoda)

The aim of this study was to evaluate the levels of phylogenetic heritability of the geographical range size, shape and position for 88 species of fiddler crabs of the world, using phylogenetic comparative methods and simulation procedures to evaluate their fit to the neutral model of Brownian motion. The geographical range maps were compiled from literature, and range size was based on the entire length of coastline occupied by each species, and the position of each range was calculated as its latitudinal and longitudinal midpoint. The range shape of each species was based in fractal dimension (box‐counting technique). The evolutionary patterns in the geographical range metrics were explored by phylogenetic correlograms using Moran’s I autocorrelation coefficients, autoregressive method (ARM) and phylogenetic eigenvector regression (PVR). The correlograms were compared with those obtained by simulations of Brownian motion processes across phylogenies. The distribution of geographical range size of fiddler crabs is right‐skewed and weak phylogenetic autocorrelation was observed. On the other hand, there was a strong phylogenetic pattern in the position of the range (mainly along longitudinal axis). Indeed, the ARM and PVR evidenced, respectively, that ca. 86% and 91% of the longitudinal midpoint could be explained by phylogenetic relationships among the species. The strong longitudinal phylogenetic pattern may be due to vicariant allopatric speciation and geographically structured cladogenesis in the group. The traits analysed (geographical range size and position) did not follow a Brownian motion process, thus suggesting that both adaptive ecological and evolutionary processes must be invoked to explain their dynamics, not following a simple neutral inheritance in the fiddler‐crab evolution.     

Taxonomy and biogeography of the fiddler crabs (Ocypodidae: Genus Uca) of the Atlantic and Gulf coasts of eastern North America

Fifteen species of fiddler crabs are reported for eastern North America between Massachusetts and Quintana Roo, Mexico. Thirteen occur in the United States and 11 in Mexico, with eight in common to the two countries. Of 13 species in the Gulf of Mexico, five are endemic and a sixth is restricted largely to the peninsulas of Florida and Yucatan. The status of U. rapax in the northern Gulf remains to be resolved. Range limits of most species approximate one or the other of two sets of intersecting thermal and geological boundaries that subdivide the Gulf of Mexico along north-south and east-west axes. Species belonging to subgenus Minuca tend to replace one another at the thermally-controlled Carolinian-Caribbean marine biotic boundary across the Florida peninsula and northern Gulf. However, only U. minax of all the North American fiddler crabs exhibits the classical disjunct Carolinian distribution, and this appears basically to reflect the discontinuous distribution of temperate salt marshes that are the habitat of the species. Distributions of species belonging to subgenus Celuca adhere for the most part to the subdivision of the Gulf into western terrigenous and eastern carbonate sedimentary provinces. The northern transition occurs in the vicinity of Apalachee Bay and the southern at Laguna de Terminos. A third distributional pattern is shown by U. subcylindrica , a specialized endemic species of the hypersaline Laguna Madre system of the western Gulf. The level of endemism in the fiddler crabs is relatively high in comparison with that of other marine groups within the Gulf of Mexico. This may be a consequence of the adaptations of fiddler crabs as specialized deposit feeders to regional differences in climatic and edaphic characteristics of a marginally marine upper shore habitat. The distributional patterns of the endemics could prove useful in reconstructing palaeoecological events of evolutionary significance within the Gulf of Mexico.