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.     

Bilateral linkage of monomorphic and dimorphic limb sizes in fiddler crabs

In this study, the subject of whether investment in one bilateral structure was linked to investment in the homologous bilateral opposite structure was investigated. Male fiddler crabs (genus Uca, family Ocypodidae) displayed strong bilateral claw differentiation of function and size, which are used for feeding (minor claw) or display/combat (major claw). Females had similar‐sized feeding claws. Linkage between claw size was investigated by estimating the deviations from an overall fitted regression of claw length to body size. The positive correlations of the deviations of claw size for major and minor claws of males and between right and left claws of females, relative to body size, suggested a linkage in investment between one claw and the corresponding claw on the other side of the body, for both monomorphic females and dimorphic males. A signal to send resources may be effectively gated to the claw complex, suggesting that positively correlated resources are allocated to both claws. Positive correlations were also found at the interspecific level. The fiddler crab model, described here, gives access to study the linkage in symmetric and asymmetric bilateral structures in the same species with a connection to the macroevolutionary level.     

Prey-induced changes to a predator's behaviour and morphology: Implications for shell-claw covariance in the northwest Atlantic

Whereas many plasticity studies demonstrate the importance of inducible defences among prey, far fewer investigate the potential role of inducible offences among predators. Here we ask if natural differences in a snail's shell hardness can induce developmental changes to a predatory crab's claw size. To do this, we fed Littorina obtusata snails from either thick- or thin-shelled populations to captive European green crabs Carcinus maenas. The crabs' shell-breaking behaviour dominated among those fed thin-shelled snails, whereas crabs fed thick-shelled snails mostly winkled flesh through the shell opening without damaging the shell itself (a.k.a. aperture-probing behaviour). Significantly, the size of crab crusher claws grew in proportion to the frequency of shell-crushing behaviour and, for a same shell-crushing frequency, crabs fed thick-shelled snails grew larger crusher claws than those fed thin-shelled snails after two experimental moults. Diet and behaviour had no effect on the growth of the smaller cutter claws of same individuals, providing good evidence that allometric changes to crusher claws were indeed a result of differential use while feeding. Findings indicate that both predation habits and claw sizes are affected by green crabs' diet, supporting the hypothesis that prey-induced phenotypic plasticity contributes to earlier accounts of shell-claw covariance between this predator and its Littorina prey in the wild.     

Specialized shell-breaking crab claws in Cretaceous seas

Here we report on a large brachyuran crab species from the Late Cretaceous of Mexico that has claws indicative of highly specialized shell-breaking behaviour. This crab possessed dimorphic claws (the right larger than the left), armed with several broad teeth, including a curved tooth structure found at the base of the movable finger of the right claw. The curved tooth is similar to the one observed on claws of many living durophagous crabs that use it as a weapon to peel, crush or chip the edges of hard-shelled prey, particularly molluscs. These morphological traits suggest that specialized shell-breaking crab predators had evolved during the Cretaceous, which contradicts previous findings supporting an Early Cenozoic origin for specialized shell crushers within the brachyuran clade.     

Sex, size and colour in a semi-terrestrial crab, Heloecius cordiformis (H. Milne Edwards, 1837)

We investigated the relationship between sex, size and colour in the little studied Australian endemic semaphore crab, Heloecius cordiformis, and related it to the crabs' social system with the aim of identifying the potential signalling function of claw colour.Equal sampling of crabs from all size classes revealed a strong relationship between sex, size and claw colour. Purple-clawed males were larger and had larger claws than pink-, orange- or green-clawed males. Male claws showed positive allometric growth: relative to body size, purple-clawed males had larger claws. The largest females had pink claws; the few with purple claws were no larger than immature green-clawed crabs. Female claws grow isometrically with the body so the relative claw size did not differ among the female colour classes. Quantitative measurements of claw colour revealed spectral differences between these subjectively described colours. The purple claws typical of large males also contrasted more strongly against the mudflat background than the other colours.Heloecius copulate outside female-owned burrows and probably within male-owned burrows. The male's waving display, in which both claws are raised and lowered, may feature in both mating strategies: as a territorial display and to attract wandering females. Large males are competitively superior so size, and potentially colour, are important in territorial disputes and may also feature in mate choice.     

The Effects of a Competitor on the Foraging Behaviour of the Shore Crab Carcinus maenas

Optimal Diet Theory suggests that individuals make foraging decisions that maximise net energy intake. Many studies provide qualitative support for this, but factors such as digestive constraints, learning, predation-risk and competition can influence foraging behaviour and lead to departures from quantitative predictions. We examined the effects of intraspecific competition within a classic model of optimal diet – the common shore crab, Carcinus maenas, feeding on the mussel, Mytilus edulis. Unexpectedly, we found that breaking time (Tb), eating time (Te), and handling time (Th) all decreased significantly in the presence of a conspecific. Reduced handling time in the presence of a competitor resulted in an increased rate of energy intake, raising the question of why crabs do not always feed in such a way. We suggest that the costs of decreased shell breaking time may be increased risk of claw damage and that crabs may be trading-off the potential loss of food to a competitor with the potential to damage their claw whilst breaking the shell more rapidly. It is well documented that prey-size selection by crabs is influenced by both the risk of claw damage and competition. However, our results are the first to demonstrate similar effects on prey handling times. We suggest that crabs maximise their long-term rate of energy intake at a scale far greater than individual foraging events and that in order to minimise claw damage, they typically break shells at a rate below their maximum. In the presence of a competitor, crabs appear to become more risk-prone and handle their food more rapidly, minimising the risk of kleptoparasitism.     

The geography of crushing: Variation in claw performance of the invasive crab Carcinus maenas

The major claws of predatory, durophagous decapods are specialized structures that are routinely used to crush the armor of their prey. This task requires the generation of extremely strong forces, among the strongest forces measured for any animal in any activity. Laboratory studies have shown that claw strength in crabs can respond plastically to, and thereby potentially match, the strength of their prey's defensive armor. These results suggest that claw strength may be variable among natural populations of crabs. However, very few studies have investigated spatial variation in claw strength and related morphometric traits in crabs. Using three geographically separate populations of the invasive green crab in the Gulf of Maine, we demonstrate, for the first time, geographic variation in directly measured claw crushing forces in a brachyuran. Despite variation in mean claw strength however, the scaling of claw crushing force with claw size was consistent among populations. We found that measurements of crushing force were obtained with low error and were highly repeatable for individual crabs. We also show that claw mass, independent of a linear measure of claw size, and carapace color, which is an indicator of time spent in the intermoult, were important predictors of claw crushing force.     

The influence of temperature,sex and chela size in the foraging strategy of the shore crab,Carcinus maenas (L.)

Foraging rate was highly variable among shore crabs of the same size category and for individual crabs from day to day. Possible physiological reasons for this variability are discussed. Shore crab foraging rate, both in terms of mussels eaten per day and energy intake per day, was estimated to be higher at 17°C than at 10°C. The shape of diet curves and their mode for male shore crabs at 17°C closely resembled those for 10°C, indicating that the temperature increase had no effect on their previously demonstrated optimal foraging strategy.

Female and certain male shore crabs showed a preference for prey smaller than for other equivalent sized males. These suboptimally feeding male and female crabs attained a relatively higher prédation rate (mussels day^‐1), although their energy intake (KJ day^‐1) remained lower than that of optimally feeding males. Preferred mussel size, number of mussels eaten per day and energy intake were strongly related to master chela height. The diet curves for female and suboptimally feeding male shore crabs could be explained by these crabs’ proportionately smaller master chelae.     

Flow visualisation and high speed video analysis of water jets in the snapping shrimp (Alpheus heterochaelis)

Snapping shrimp (Alpheus heterochaelis) produce a fast, well-focused water jet by rapid closure of their specialised snapper claw. As shown previously, water jets may injure the opponent in interspecific encounters (e.g. with small crabs) although no damage was observed in intraspecific encounters. For conspecific receivers the jet represents a potential hydrodynamic signal and can be analysed with the help of mechanosensory hairs. To gain more insight in the biophysical characteristics of the water jet we visualised and analysed jets of tethered snapping shrimp using standard and high speed video recordings. Water jet width increases with increasing distance from the snapper claw tip, and both width and distance increase with increasing snapper claw size. Water jet distances do not increase with increasing claw cocking duration (building up muscle tension) but medium cocking durations of about 550 ms result in longest distances. Mean water jet velocity is 6.5 m s⁻¹ shortly after claw closure but rapidly decreases subsequently. At the mean distance between snapping conspecifics (9 mm) water jet velocities produced by snapping shrimp with larger snapper claws are significantly higher than those of animals with smaller claws. Interestingly, males with equal snapper claw size as females produce significantly faster water jets. Accepted: 31 March 1999     

Regulation of sex-specific feeding behavior in fiddler crabs: physiological properties of chemoreceptor neurons in claws and legs of males and females

This study examined properties of chemoreceptor neurons in the claws and legs of the fiddler crabs Uca pugilator and U. pugnax. The primary goal was to establish the neural basis of previously observed greater female sensitivity to feeding stimulants, and secondarily to compare physiological properties of chemoreceptor neurons in these semi-terrestrial crustaceans with those of fully aquatic forms. Sensitivity of chemoreceptor neurons in claws and legs is sex-specific; individual neurons of females respond to lower stimulus concentrations than male chemoreceptor neurons, and equivalent concentrations elicit greater spiking in female vs male chemoreceptor neurons. Thus, the population of chemoreceptor neurons in females expresses lower thresholds and greater average sensitivity than in males. Greater sensitivity of claw neurons explains observations indicating that females continue to feed at food levels too low to stimulate males. Sensitivity differences in leg neurons of males vs females have no clear behavioral correlate, but suggest that females can orient to more dilute stimuli than males. Chemoreceptor neurons of fiddler crabs have low sensitivities and slow rates of adaptation compared to other crustaceans. Also, neurons in claws adapt less slowly than neurons in legs, which may reflect subtle differences in the chemical stimulus environment experienced by claws vs legs.     

The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

Male mating success in a fiddler crab: a lesson in sample sizes

Autotomy and regrowth of a body part occurs in many animal species. It is costly to regrow the limb and there are often additional long-term costs in, for example, limb strength, foraging efficiency and even mating success. In the fiddler crab Uca mjoebergi, 7 % of males have autotomized and regrown their large claw at some point in their lives. Previous work has shown that there is a great disadvantage to having a regenerated claw. While these males are able to attract mate-searching females to visit them, none of the 84 males observed to have mated in previously collected data had regenerated claws. Since females’ final mate choice is based on burrow structure, it was assumed that males with regenerated claws had poorer burrows. Here we show that, by finding only three cases of a female mating with a regenerated claw male, that there is, in fact, no mating disadvantage to having a regenerated claw. We also show that the burrows of males with regenerated claws are no different than those of orginal-clawed males. This is a very clear reminder that sample size matters, especially when dealing with rare events.     

Lameness and Claw Lesions of the Norwegian Red Dairy Cattle Housed in Free Stalls in Relation to Environment,Parity and Stage of Lactation

Approximately 88% of Norwegian dairy cattle are housed in tie stalls. Free stall housing for all dairy cattle will be implemented within 20 years. This means that the majority of existing stalls will be rebuilt in the near future. Fifty-seven free stall herds of the Norwegian Red breed were randomly selected and 1547 cows and 403 heifers were trimmed by 13 claw trimmers during the late winter and spring of 2002. The claw trimmers had been taught diagnosing and recording of claw lesions. Environment, management- and feeding routines were also recorded. Fifty-three herds had concrete slatted alleys while 4 had solid concrete. Thirty-five herds had concrete as a stall base, while 17 had rubber mats, 2 had wood and 3 had deep litter straw beds. The prevalence of lameness was 1.6% in hind claws. Models for lameness and claw lesions were designed to estimate the influence of different risk factors and to account for the cluster effects within herd and claw trimmer. Detected risk factors for lameness were: parity three and above and narrow cubicles; for heel horn erosions: lactation stage around 5–7 months after calving and solid concrete alleys; for haemorrhages of the white line: lactation stage around 3–5 months after calving and solid concrete alleys; for haemorrhages of the sole: parity one, lactation stage around 5–7 months after calving and short cubicles, for white line fissures: slatted concrete alleys; for asymmetrical claws: parities two and above and for corkscrewed claws: solid concrete alleys. The prevalence of lameness in heifers was low, however 29% had one or more claw lesions. Heifers that were housed in pens or free stalls had more heel-horn erosions, haemorrhages of the sole and white-line fissures than heifers in tie stalls. As new free stalls are being built, it is important to optimise the conditions for claw health.     

Even Weak Males Help Their Neighbours: Defence Coalitions in a Fiddler Crab

Large male fiddler crabs sometimes help smaller neighbours to defend their territories against intruders. These coalitions occur when the helper is likely to defeat the intruder (helper larger than intruder) and the neighbour is likely to lose his territory without help (intruder larger than neighbour). Previous studies of coalitions have excluded males with regenerated claws. Such claws are weaker weapons that make the bearer competitively inferior. Here, we show that male Uca annulipes with regenerated claws are as likely as males with original claws to help their neighbours in territory defence, even though, as weaker males they potentially pay greater costs, being more likely to lose their undefended burrow. We suggest that males with regenerated claws gain greater benefits from retaining a current, small neighbour and that, as in non‐coalition fights, the regenerated claw acts as a visual bluff in the early stages of combat. Furthermore, we show that intruders with regenerated or original claws are equally likely to be attacked by a ‘helping’ neighbour. This bolsters the argument that males cannot visually differentiate between original and regenerated claws.     

Relationship of claw form and exoskeleton condition to reproductive system size and methyl farnesoate in the male spider crab,Libinia emarginata

Summary

Methyl farnesoate (MF) expression and reproductive system size were compared in five representative groups of male L. emarginata selected from a sample collected in November. The groups differed from each other with respect to carapace size (small, intermediate and large), relative propodus size (small and large claw forms), and condition of the exoskeleton (abraded and unabraded). Large males with large claws and abraded exoskeletons had reproductive system indices which were significantly larger than any other group. The mandibular organs of these crabs also had significantly higher rates of methyl farnesoate synthesis in vitro. Hemolymph titers of methyl farnesoate were also highest in this group, but were not significantly different from the group with small carapaces, small claws and unabraded exoskeletons. Methyl farnesoate titers were significantly lower in all other groups of unabraded animals with small or large claws. These results suggest that methyl farnesoate may play a role in morphogenesis and reproduction in male L. emarginata.     

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.     

Mud crabs (Xanthidae) in Chesapeake Bay: claw characteristics and predation on epifaunal bivalves

Abstract. We examined claw characteristics of mud crabs (Eurypanopeus depressus, Rhithropanopeus harrisii) to determine if one crab species was potentially more powerful than the other. We related our findings to the abilities of individuals of each species to open epifaunal mytiliform bivalves (Ischadium recurvum; Mytilopsis leucophaeata) that occur on beds of eastern oysters (Crassostrea virginica) in mesohaline Chesapeake Bay. There were high correlations between claw width or height and claw length, and between claw length and carapace width for both mud crab species. The mechanical advantage or “grip strength’ of the crusher and cutter claws of both species did not change with crab size (carapace width) and did not differ between sexes in each species, nor did the cutter data differ between species. However, individuals of E. depressus had a significantly stronger crusher claw grip than did those of R. harrisii. Data on mechanical advantage for both species were similar to values reported in the literature for members of other xanthid crab species. These values in turn overlapped those reported for calappid, cancrid, majid, and grapsid crabs, and were greater than those of various species of portunid crabs and individual species of fiddler crab, lobster, crayfish, and ghost shrimp. When simultaneously presented with the two species of bivalves, the mud crabs E. depressus chose mussels of M. leucophaeata first and crabs of R. harrisii chose mussels of I. recurvum first about two‐thirds of the time; ultimately, the crabs ate both bivalve species in >50% of the choice experiments. The size range in E. depressus was greater than that in R. harrisii, and crabs of E. depressus opened larger bivalves than did crabs of R. harrisii, although similar‐sized individuals of the two crab species overlapped in their ability to open bivalves of both species. In Mytilopsis leucophaeata, there is probably no size refuge from predation by the mud crabs whereas the larger mussels of I. recurvum do have a refuge in size.     

Maximum force production: why are crabs so strong?

Durophagous crabs successfully hunt hard-shelled prey by subjecting them to extremely strong biting forces using their claws. Here I show that, for a given body mass, six species of Cancer crabs (Cancer antennarius, Cancer branneri, Cancer gracilis, Cancer magister, Cancer oregonensis and Cancer productus) were able to exert mean maximum biting forces greater than the forces exerted in any other activity by most other animals. These strong biting forces were in part a result of the high stresses (740-1350 kN m(-2)) generated by the claw closer muscle. Furthermore, the maximum muscle stress increased with increasing mean resting sarcomere length (10-18 microm) for the closer muscle of the claws of these six Cancer species. A more extensive analysis incorporating published data on muscle stresses in other animal groups revealed that stress scales isometrically with the resting sarcomere length among species, as predicted by the sliding filament model of muscle contraction. Therefore, muscle or filament traits other than a very long mean sarcomere length need not be invoked in explaining the high stresses generated by crustacean claws.     

It takes two: Seasonal variation in sexually dimorphic weaponry results from divergent changes in males and females

Sexually dimorphic weaponry often results from intrasexual selection, and weapon size can vary seasonally when costs of bearing the weapon exceed the benefits outside of the reproductive season. Weapons can also be favored in competition over nonreproductive resources such as food or shelter, and if such nonreproductive competition occurs year‐round, weapons may be less likely to vary seasonally. In snapping shrimp (Alpheus angulosus), both sexes have an enlarged snapping claw (a potentially deadly weapon), and males of many species have larger claws than females, although females are more aggressive. This contrasting sexual dimorphism (larger weaponry in males, higher aggression in females) raises the question of whether weaponry and aggression are favored by the same mechanisms in males and females. We used field data to determine whether either sex shows seasonal variation in claw size such as described above. We found sexual dimorphism increased during the reproductive season due to opposing changes in both male and female claw size. Males had larger claws during the reproductive season than during the nonreproductive season, a pattern consistent with sexual selection. Females, however, had larger claws during the nonreproductive season than during the reproductive season—a previously unknown pattern of variation in weapon size. The observed changes in female weapon size suggest a trade‐off between claw growth and reproduction in the reproductive season, with investment in claw growth primarily in the nonreproductive season. Sexually dimorphic weaponry in snapping shrimp, then, varies seasonally due to sex differences in seasonal patterns of investment in claw growth, suggesting claws may be advantageous for both sexes but in different contexts. Thus, understanding sexual dimorphisms through the lens of one sex yields an incomplete understanding of the factors favoring their evolution.