Depth cues, behavioural context, and natural illumination: some potential limitations of video playback techniques

By expanding on issues raised by D’Eath (1998), I address in this article three aspects of vision that are difficult to reproduce in the video- and computer-generated images used in experiments, in which images of conspecifics or of predators are replayed to animals. The lack of depth cues derived from binocular stereopsis, from accommodation, and from motion parallax may be one of the reasons why animals do not respond to video displays in the same way as they do to real conspecifics or to predators. Part of the problem is the difficulty of reproducing the closed-loop nature of natural vision in video playback experiments. Every movement an animal makes has consequences for the pattern of stimulation on its retina and this ”optic flow” in turn carries information about both the animal’s own movement and about the three-dimensional structure of the environment. A further critical issue is the behavioural context that often determines what animals attend to but that may be difficult to induce or reproduce in an experimental setting. I illustrate this point by describing some visual behaviours in fiddler crabs, in which social and spatial context define which part of the visual field a crab attends to and which visual information is used to guide behaviour. I finally mention some aspects of natural illumination that may influence how animals perceive an object or a scene: shadows, specular reflections, and polarisation reflections. Received: 23 November 1999 / Received in revised form: 9 February 2000 / Accepted: 10 February 2000     

Behavioural evidence for body colour signaling in the fiddler crab Uca perplexa (Brachyura: Ocypodidae)

In the reproductive season, mature females of the fiddler crab Uca perplexa leave their burrows and wander about their habitat for mating. To clarify whether the fiddler crabs respond to colour or luminosity, I examined the behavioural responses of the males to the wandering females before and after the females were painted white, red, black or blue. The behaviours of the males were categorized into three types: lateral–circular wave and lateral–straight wave for courting, and repelling. Before painting, almost all of the males courted the females. After painting, significantly fewer males courted the red-, black- and blue-painted females than courted the white-painted females. These results mean that the fiddler crabs can discriminate colours or luminosity. The role of body colour as a visual signal in crab society is discussed.     

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.     

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.     

A Spatially Explicit Model of Synchronization in Fiddler Crab Waving Displays

Fiddler crabs (Uca spp., Decapoda: Ocypodidae) are commonly found forming large aggregations in intertidal zones, where they perform rhythmic waving displays with their greatly enlarged claws. While performing these displays, fiddler crabs often synchronize their behavior with neighboring males, forming the only known synchronized visual courtship displays involving reflected light and moving body parts. Despite being one of the most conspicuous aspects of fiddler crab behavior, little is known about the mechanisms underlying synchronization of male displays. In this study we develop a spatially explicit model of fiddler crab waving displays using coupled logistic map equations. We explored two alternative models in which males either direct their attention at random angles or preferentially toward neighbors. Our results indicate that synchronization is possible over a fairly large region of parameter space. Moreover, our model was capable of generating local synchronization neighborhoods, as commonly observed in fiddler crabs under natural conditions.     

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.     

Invertebrate perception: measuring depth intervals through path integration and vision

A foraging fiddler crab can estimate how close a potential intruder is from its burrow entrance, even when the entrance in the sand is invisible to the crab. Recent work shows that, to assess this depth interval, crabs combine information from vision and path integration in an unusual manner.     

Habitat partitioning between prey soldier crab Mictyris brevidactylus and predator fiddler crab Uca perplexa

Interaction and habitat partition between the soldier crab Mictyris brevidactylus (prey) and the fiddler crab Uca perplexa (predator) were examined at a sandy tidal flat on Okinawa Island, Japan, where they co-occur. Both live in dense colonies. When the soldier crabs were released in the densely populated habitat of the fiddler crab, male fiddler crabs, which maintain permanent burrows in hard sediment, preyed on small soldier crabs and repelled large ones. Thus, the fiddler crabs prevented the soldier crabs from trespassing. It was also observed whether soldier crabs burrowed successfully when they were released 1) where soldier crab burrows just under the sand were abundant, 2) in a transition area between the two species, 3) an area without either species, and 4) where artificial tunnels simulated soldier crabs' feeding tunnels were made by piling up sand in the area lacking either species. In contrast to the non-habitat area, many soldier crabs burrowed in the sediment near the release point in the tunnel, transition and artificial tunnel areas. This indicates that the feeding tunnels on the surface attracted other crabs after emergence. When the large male fiddler crabs were transplanted into the artificial burrows made in soft sediment of the soldier crab habitat, all left their artificial burrows by 2 days. In the fiddler crab habitat, however, about one-third of the transplanted male fiddler crabs remained in the artificial burrows after 3 days. The soldier crabs regularly disturb the sediment by the up and down movement of their burrow (small air chamber) between tides. This disturbance probably prevents the fiddler crab from making and occupying permanent burrows. Thus, it appears that these crabs divide the sandy intertidal zone by sediment hardness and exclude each other by different means.     

Colour spaces in ecology and evolutionary biology

The recognition that animals sense the world in a different way than we do has unlocked important lines of research in ecology and evolutionary biology. In practice, the subjective study of natural stimuli has been permitted by perceptual spaces, which are graphical models of how stimuli are perceived by a given animal. Because colour vision is arguably the best‐known sensory modality in most animals, a diversity of colour spaces are now available to visual ecologists, ranging from generalist and basic models allowing rough but robust predictions on colour perception, to species‐specific, more complex models giving accurate but context‐dependent predictions. Selecting among these models is most often influenced by historical contingencies that have associated models to specific questions and organisms; however, these associations are not always optimal. The aim of this review is to provide visual ecologists with a critical perspective on how models of colour space are built, how well they perform and where their main limitations are with regard to their most frequent uses in ecology and evolutionary biology. We propose a classification of models based on their complexity, defined as whether and how they model the mechanisms of chromatic adaptation and receptor opponency, the nonlinear association between the stimulus and its perception, and whether or not models have been fitted to experimental data. Then, we review the effect of modelling these mechanisms on predictions of colour detection and discrimination, colour conspicuousness, colour diversity and diversification, and for comparing the perception of colour traits between distinct perceivers. While a few rules emerge (e.g. opponent log–linear models should be preferred when analysing very distinct colours), in general model parameters still have poorly known effects. Colour spaces have nonetheless permitted significant advances in ecology and evolutionary biology, and more progress is expected if ecologists compare results between models and perform behavioural experiments more routinely. Such an approach would further contribute to a better understanding of colour vision and its links to the behavioural ecology of animals. While visual ecology is essentially a transfer of knowledge from visual sciences to evolutionary ecology, we hope that the discipline will benefit both fields more evenly in the future.     

The effect of audience on intrasexual interaction in the male fiddler crab, <Emphasis Type="Italic">Uca maracoani</Emphasis> (Latreille 1802–1803, Decapoda,Ocypodidae)

Animal communication occurring in wide networks can involve signals sent to several receivers. The animals composing the audience may affect how individuals display during an interaction and may change the message that is sent. In this study, we investigated whether the presence of a conspecific affected the intensity of agonistic interaction between male fiddler crabs, Uca maracoani. Pairs of males of the same size were observed when in the presence of a male, a female or no crab as audience. We found that if there is a female audience, males became more aggressive and interacted the most. Also, the female audience leads to an increase in incidence of male foaming, possibly indicating predisposition for mating. If the audience was a male or if there was no audience, there was no significant difference in interaction. These results indicate that the presence of an audience affects the way male fiddler crabs interact and the type of displays they show. Therefore, the context seems to guide the fiddler crab behavior in terms of how they perform in order to send information about themselves to conspecifics.     

Variability of a dynamic visual signal: the fiddler crab claw-waving display

Fiddler crabs use elaborate, species-specific claw-waving displays to communicate with rivals and mates. However, detailed comparative studies of fiddler crab signal structure and structural variations are lacking. This paper provides an analysis of the claw-waving displays of seven Australian species of fiddler crab, Uca mjoebergi, U. perplexa, U. polita, U. seismella, U. signata, U. elegans and U. vomeris. We used digital video to record and analyse the fine-scale spatiotemporal properties of these movement-based visual signals. We found that the structure and timing of the displays is species-specific, exhibiting inter-specific differences that follow phylogenetic relationships. The displays showed intra-specific variation according to individual identity, geographic location and fine-scale behavioural context. The observed differences and variations are discussed in the light of the evolutionary forces that may shape their design.     

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.     

The blue streak: a dynamic trait in the mud fiddler crab, Uca pugnax

Mud fiddler crabs, Uca pugnax, have a streak of blue coloration located on the front of the carapace above the mouth and centered between the eyes. We documented that this blue streak is absent in juveniles and develops as crabs become sexually mature. By photographing male crabs under controlled conditions in the laboratory, we demonstrated that the brightness of the blue streak (in comparison with the rest of the carapace) is dynamic, and can dim from bright blue to nearly black in fewer than two minutes. We examined blue streak variability in male crabs in response to physical factors (light and temperature) and social context to begin to understand what causes its dynamic response. The blue streak darkens in response to decreased ambient light, but does not respond to changes in temperature. In the field, it is brighter when crabs are roaming on the mudflat or fighting, but darker when crabs are basking or performing waving displays. The highly visual nature of fiddler crabs and the dynamic character of the blue streak suggest that it may communicate information about the state of a crab or its environment.     

Comparative Studies of Reproductive Behavior in Mantis Shrimps and Fiddler Crabs

Comparative ethological studies of closely related species canplace into clear perspective the functions of behavioral traitsas species diverge and adapt to different environments. In thispaper we contrast the reproductive behavior of species withintwo crustacean groups: the mantis shrimps (stomatopods) andthe fiddler crabs (genus Uca). For the stomatopods, we identifyprobable selective relationships between features of species'ecology and their mating systems. Population density, the kindand availability of shelters in which these shrimps live, theintensity of predation and spawning cycles all play importantroles in molding reproductive behavior in this group. Associationsbetween the ecology and mating systems of fiddler crabs havebeen discussed recently elsewhere. Here we focus on a comparativeanalysis of sexual communication in these crabs. A study ofcompetitive courtship signaling and mate choice in the fiddlercrab Uca beebei has shown that certain male courtship signalsare highly attractive probably because they exploit female sensory-responsesystems that have been molded by selection for escape from predators.Interspecific comparison of male courtship displays and theresponses of females to these displays suggest that sensoryexploitation may play an important role in the evolution ofsexual signals in the genus. Comparative studies have advancedour understanding of how natural and sexual selection affectthe reproductive behavior of both stomatopods and fiddler crabs.     

The relative importance of substratum characteristics and recruitment in determining the spatial distribution of the fiddler crab Uca uruguayensis Nobili

The roles of sediment characteristics and the pattern of recruitment in influencing the abundance of the fiddler crab Uca uruguayensis on Argentinean mudflats were evaluated. The density of adult crabs showed a patchy distribution related to the sediment thickness (depth at which a layer of fossil shells are buried), but the density of juvenile crabs was not coupled with the density of adult crabs. In a field experiment, fossil shells were removed and the density of crabs significantly increased, which demonstrates that the presence of the layer of shells is a structure that may hinder the establishment of burrows. The density of crabs was related to sediment thickness, sediment torque and organic matter content. The importance of each of these variables was different for adult and juvenile crabs, indicating that the distribution of adult crabs may be caused by mechanisms affecting adult crabs themselves and is not established by the recruitment pattern. Moreover, in a field experiment, the density of juveniles decreased when adult crabs were added, and increased when adult crabs were removed.The morphology of burrows was related to sediment characteristics. Burrows were deeper, longer and more voluminous when sediment thickness was high. The volume of burrows decreased with increasing sediment torque. These results suggest that the morphology of burrows is related to the space available and the ease with which sediment it can be excavated. However, an important amount of variability remained unexplained, suggesting the presence of additional environmental variables or behavioural plasticity not considered by this study. Together, these results demonstrate that the spatial heterogeneity in the environmental factors can be translated to a spatial heterogeneity in the distribution of fiddler crabs.     

A remarkable case of fiddler crab, Uca spp., alpha diversity in Wallacea

A preliminary attempt was made to quantify the alpha diversity of fiddler crabs (Uca spp.) along a short transect across the interface between an intertidal mudflat and a largely Avicennia mangrove at Ambeua on the Indonesian island of Kaledupa in Sulawesi Tenggara, by means of visual counts within 2 m² quadrats. An astounding diversity of up to six seemingly coexisting species per quadrat—and a total of seven such species within an area of 4 m²—was revealed. Four species of the ecologically equivalent sentinel crabs (Macrophthalmus) were also present in the quadrats. Seven fiddler crab species equates to more than half those known from Wallacea, more than one-third of all those known from Indonesia, and almost one-fifth of the entire Uca fauna of the Indo-West-Pacific.     

Stimulation of feeding behavior in three species of fiddler crabs by hexose sugars

Feeding responses to five hexoses were examined in three closelyrelated species of fiddler crabs, Uca minax, U.pugnax and U.pugilator.Hexoses tested were glucose, galactose, sorbose, fructose andmannose. Intact crabs and eyestalk-ablated crabs were tested.Responses to sugars were species specific. Eyestalks are directlyinvolved in vision and overall neural integration as well aswith chemosensory and metabolic pathways associated with feeding.Overall, eyestalk-ablated crabs were more sensitive uian intactcrabs. Studies of responses of individuals within a populationto hexoses showed tiiere are individual U.pugnax that respondto galactose and others that do not. Similarly, there were U.pugilatorindividuals that were mannose responsive and others that didnot respond to mannose. An additional study of differences inpopulation responses to hexoses would provide valuable toolsin studying geographic relationships between fiddler crab populations.     

Periphyton density and shading in relation to tidal depth and fiddler crab activity in intertidal seagrass beds of the Banc d'Arguin (Mauritania)

Periphyton development was studied on microscopic glass slides and leaves of Zostera noltii Hornem. in an intertidal area in the Banc d'Arguin (Mauritania). The effects of shading, tidal depth and grazing activities by the fiddler crab Uca tangeri Eydoux were evaluated. For all experiments, periphyton ash content was high (52–93%) and ash-free dry weight ranged between 0.10–0.63 mg cm^–2. Slides accumulated more periphyton than leaves.Artificial shading (62–99%) for 13 days had no effect on periphyton densities on leaves. Increased tidal depth resulted in higher ash-free dry weight on slides, but in lower ash-free dry weight on leaves. Significant variation along the coastline also existed. The effect of fiddler crabs was studied using exclosures. Presence of fiddler crabs reduced periphyton density on slides, whereas light transmittance was increased. On leaves, no significant fiddler crab effect was found. This difference between leaves and slides was probably caused by a storm at the day before the end of the experiment, and by the higher periphyton density on slides as compared with leaves. As visual inspection during the experiment showed clear differences in appearance of leaves inside and outside the exclosures, the storm probably sloughed off mainly the older leaves, i.e. those on which the differences in periphyton cover were the highest.It is hypothesized that periphyton accumulation is higher with increased tidal depth, whereas fiddler crab grazing pressure also increases in this direction. The result is a decreased periphyton density with increased tidal depth.The presently found light extinction coefficients (mean 0.8 m^–1) and periphyton light attenuance (up to 25%) in Banc d'Arguin are not likely to affect seagrass leaf growth.     

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.     

Effect of cadmium exposure on lactate dehydrogenase activity in the hepatopancreas and abdominal muscle of the fiddler crab,Uca pugilator

1. Lactate dehydrogenase (LDH) activity in the hepatopancreas and abdominal muscle of fiddler crabs, Uca pugilator, was determined after 24 and 48 hr of exposure to 2 ppm cadmium chloride.2. For the cadmium exposed crabs, LDH activity in the hepatopancreas decreased, whereas that in the abdominal muscle increased.3. The increased LDH activity in the abdominal muscle may reflect increased dependence on anaerobic carbohydrate metabolism in fiddler crabs exposed to cadmium in their environment.