Force, function and mechanical advantage in the chelae of the American lobster Homarus americanus (Decapoda: Crustacea)

The dimorphic chelae from both sexes and a wide size range of American lobsters, Homarus americanus , were studied with respect to allometry, mechanical advantage, closer muscle apodeme area and occlusive surface morphology. The maximum forces produced by the crusher and cutter chelae were estimated by an in vitro and a static in vivo technique. Another in vivo technique, involving strain gauges, was used to measure the forces delivered by crusher chelae. The latter technique gave data on force pulse duration and frequency, and in combination with a video-recording system could be useful for future studies of predation behaviour. The maximum forces generated increased with chela height for both crusher and cutter chelae. A maximum force of 256 Newtons (N) was recorded near the middle of the crusher dactyl from a 172-mm carapace length lobster, by the strain gauge technique. Crusher chelae developed larger maximum forces than cutter chelae of the same height. This was attributable more to the crusher chela's higher mechanical advantage than to its developing higher input forces. The mean mechanical advantage for male crusher (0.33) was significantly higher than that for female crusher chelae (0.29). Male and female cutter chelae had the same mean mechanical advantage values (0.16). Values for maximum stress developed during contraction in both the crusher and cutter chela closer muscles decreased with chela size. The morphology of the chelae correlated to the forces produced and predation behaviour.     

Size,shape, and sex‐dependent variation in force production by crayfish chelae

The ability to generate large closing forces is important for many animals. Several studies have demonstrated that bite or pinching force capacity is usually related to the linear dimensions of the closing apparatus. However, relatively few studies have applied geometric morphometrics to examine the effects of size‐independent shape on force production, particularly in studies of crustacean pinching force. In this study, we utilized traditional and geometric morphometric techniques to compare the pinching force of Procambarus clarkii crayfish to their chela morphology. We found that males possessed larger chelae and pinched harder than females, but that their chela shape and size were weak predictors of strength. Female pinching force was significantly affected by both chela size and shape, with shape variation along the short axis of the claw contributing most to pinching force. We discuss our results in the context of reliable signaling of strength by males and females, and the different selective forces acting on chela shape in the two sexes.     

Heterochely and handedness in the shore crab Carcinus maenas (L.) (Crustacea: Brachyura)

A large sample ofadult male Carcinus maenas was 79% right-handed and 21% left-handed. A separate sample of 207 intact adult males was divided into left-handed and right-handed crabs and four measurements were taken from all major and minor chelae. Correlation and regression analyses against carapace width on log-transformed data showed that major chelae of right-handed crabs grow proportionately higher with increasing size and the ideal mechanical advantage increases; concurrently, the fingers of the minor chelae grow proportionately longer. The data for left-handed crabs showed greater variability, especially for minor chelae, providing evidence for the concept that left-handedness arises by reversal of handedness following loss of the major chela from the right-hand side. Records of handedness in large samples of non-ocypodid heterochelous brachyuran crabs show a preponderance of right-handedness.     

Structure and function of the chelae and chela closer muscles of the shore crab Carcinus maenas (Crustacea: Brachyura)

Major and minor chelae of Carcinus maenas were investigated with respect to mechanical advantage, angles of pennation and sarcomere lengths of the closer muscle fibres, maximum pinching forces induced by nervous stimulation and maximum stress in the closer muscles. Major chelae differed from minor chelae in all these factors. Mean mechanical advantages of major and minor chelae were 0·36 and 0·26, respectively. Angles of pennation were greater at the distal ends of closer muscles and declined proximally: in half open major and minor chelae distal to proximal angles were about 35–26° and 29–19°, respectively. Mean sarcomere lengths in closer muscles of fully closed major and minor chelae were significantly different at 7·6 and 6·0 fim, respectively; there was great variation within chelae and between crabs. Major chelae pinched more strongly than minor chelae due to higher mechanical advantages, larger closer apodemes and greater stress in the muscle. Mean maximum stresses in major and minor chelae were 677 and 474 kN/m², respectively. The biggest stress recorded–1057 kN rrr²–was in a major chela. Pinching forces induced by injecting a high K⁺ and caffeine ringer were about 20% weaker than nervously stimulated pinches.     

Morphology and muscle stress of chelae of temperate and tropical stone crabs Menippe mercenaria

Chela morphology and muscle stress were compared between temperate and tropical populations of stone crabs Menippe mercenuriu (Say) to test whether environmental differences might result in greater crushing strength in the tropics. Such differences include increased crab diversity in the tropics (which might lead to greater fighting among congeners), increased prey exoskeleton calcification in the tropics, and year round chela use in the tropics as opposed to seasonal chela use in the temperate population.
No latitudinal differences were found in any aspect of chela morphology, including relative chela size, mechanical advantage, apodeme surface area, and angle of muscle fibre pinnation. Summer measurements of crusher chela muscle stress were also similar between the two populations.
The maximum muscle stress determined for M. mercenariu was 220 N ∼r n -∼, much higher than stress levels previously reported for crustaceans. Other researchers have typically measured forces either from autotomized chelae or by measuring forces required to open a closed chelae. I have determined muscle stress using a force transducer that measures active gripping strength in live crabs.     

Dishonest signals of strength in male slender crayfish (Cherax dispar) during agonistic encounters

Many animals resolve disputes without combat by displaying signals of potential strength during threatening displays. Presumably, competitors use each other's displays to assess their relative strengths, and current theory predicts that these signals of strength should generally be honest. We tested this prediction by investigating the relationships among morphology, performance, and social dominance in males of the slender crayfish Cherax dispar. Crayfish routinely use their enlarged front claws (chelae) for both intimidation and fighting, making this species ideal for studying the honesty of weapon size. We evaluated five competing models relating morphological and physiological traits to dominance during paired competitive bouts. Based on the best model, larger chelae clearly resulted in greater dominance; however, chela strength had no bearing on dominance. Thus, displays of chela size were dishonest signals of strength, and the enlarged chelae of males seemingly function more for intimidation than for fighting. In addition, an analysis of the performance of isolated chela muscle showed that muscle from male crayfish produced only half the force that muscle from female crayfish produced (236.6+/-26.4 vs. 459.5+/-71.6 kN m(-2)), suggesting that males invest more in developing larger chelae than they do in producing high-quality chela muscle. From our studies of crayfish, we believe dishonest signaling could play a greater role in territorial disputes than previously imagined.     

Decapod crustacean chelipeds: an overview

The structure, growth, differentiation and function of crustacean chelipeds are reviewed. In many decapod crustaceans growth of chelae is isometric with allometry level reaching unity till the puberty moult. Afterwards the same trend continues in females, while in males there is a marked spurt in the level of allometry accompanied by a sudden increase in the relative size of chelae. Subsequently they are differentiated morphologically into crusher and cutter making them heterochelous and sexually dimorphic. Of the two, the major chela is used during agonistic encounters while the minor is used for prey capture and grooming. Various biotic and abiotic factors exert a negative effect on cheliped growth. The dimorphic growth pattern of chelae can be adversely affected by factors such as parasitic infection and substrate conditions. Display patterns of chelipeds have an important role in agonistic and aggressive interactions. Of the five pairs of pereiopods, the chelae are versatile organs of offence and defence which also make them the most vulnerable for autotomy. Regeneration of the autotomized chelipeds imposes an additional energy demand called “regeneration load” on the incumbent, altering energy allocation for somatic and/or reproductive processes. Partial withdrawal of chelae leading to incomplete exuviation is reported for the first time in the laboratory and field inMacrobrachiumspecies.     

Pinching forces in crayfish and fiddler crabs,and comparisons with the closing forces of other animals

The pinching forces of crustaceans are in many respects analogous to the biting forces of vertebrates. We examined the effects of body size and chelae size and shape, on the closing forces of the fiddler crab, Uca pugilator, and the crayfish, Procambarus clarkii. We hypothesized that the allometric relationships would be similar among species, and comparable to those reported for other decapod crustaceans. We further hypothesized that the scaling of the closing forces of crustaceans, with respect to body size and with the geometry of the pinching or biting structures, would be similar to that of vertebrates. We found that pinching forces increased with body mass, claw dimensions, and claw mass in U. pugilator, but only with claw height and claw mass in P. clarkii. Contraction time increased with body mass for both species combined, whereas contraction speed decreased. Pooled data for these and 17 other species of decapod crustacean revealed a positive correlation between the pinching force and body mass with a scaling exponent of 0.71. These data are remarkably comparable to the values on closing forces of vertebrate jaws, with the pooled data having a scaling exponent of 0.58, slightly below the value of 0.67 predicted for geometric similarity. Maximum closing forces vary tremendously among both crustaceans and animals in general, with body size and food habits being among the most important determining factors.     

A study of normal and regenerative growth in the pistol-crab,alpheus dentipes (Guèr.), with special reference to the phenomenon of chela reversal

Summary An analysis has been made of the normal growth of the thoracic appendages in the pistol-crab,Alpheus dentipes. The chelae are the only asymmetrical appendages and, within an extensive size range, the growth-coefficient of the large chela (crusher) is smaller than that of the small chela (nipper), even in the most rapidly growing dimensions of the segment containing the growth-centre.The partition coefficient for growth in certain dimensions of the chelae is not constant but is reduced progressively during the growthperiod, due (it is suggested) to the impracticability of maintaining the intensive heterogony developed during the early stages of growth.Regenerative growth of the chelae has been studied quantitatively after autotomy of one, the other, or both normal chelae, effected once only or repeatedly as new chelae appeared at moulting. New chelae are regenerated repeatedly with little or no waning of growth intensity but in continuously growing chelae, growth intensity is substantially reduced at the end of the instar of autotomy. The amount of growth is not correlated with thenumber of days available for regeneration. When both chelae regenerate simultaneously the larger one grows relatively slower than the smaller one to the end of the instar of autotomy but afterwards this condition is reversed. When the nipper alone is autotomised, growth of the crusher is inhibited and shrinkage may occur. Crusher autotomy, on the other hand, stimulates nipper growth and to an extent which varies with the dimension.The problem of chela reversal has been discussed and certain suggestions have been preferred.     

Leverage and muscle type in crab chelae (Crustacea: Brachyura)

The chelae of Cancer pagurus and Macropipus depurator were examined with respect to mechanical advantage. The closer muscles were investigated with respect to sarcomere length in the constituent fibres and to the force developed by the whole muscle during isometric contraction. Cancer chelae have a relatively high mechanical advantage, 0.329 ± 001. Cancer closer muscles contain a high proportion of fibres with long sarcomeres, mean lengths mostly falling between 12 and 15 μm, and develop a maximum stress of about 496 kN.m⁻² during contraction. These figures are typical for "slow" crustacean muscle. The chelae of M. depurator are dimorphic. In one, the strong chela, the mechanical advantage is 0.248 ± 0.066 while in the other, the fast chela, the mechanical advantage is 0.177 ± 0.006. M. depurator closer muscles contain fibres with mean sarcomere lengths mostly falling between 6 and 10 μm. The muscle develops a maximum stress of about 145 kN.m⁻² during contraction. These figures are typical of "intermediate" crustacean muscles. "Fast" muscle fibres with short sarcomeres (about 30 um) were found in the chelae of both Cancer and M. depurator but were much commoner in the latter. Thus in Cancer a high mechanical advantage is correlated with slow muscle while in M. depurator lower mechanical advantages are broadly correlated with faster muscle. Consistent correlation between mechanical advantage and muscle type in the dimorphic chelae of M. depurator , however, is lacking. No consistent regionation of fibres with similar properties was found in the muscles.     

Aspects of the biology of the big-handed crab,Heterozius rotundifrons (Decapoda: Brachyura), from Kaikoura,New Zealand

Abstract

From January 1975 to June 1976, samples of Heterozius rotundifrons A. Milne Edwards, 1867 were taken monthly from the intertidal zone at First Bay, Kaikoura Peninsula (42°25′S, 173°42′E). Small crabs (< 8 mm carapace width) were never common; they were either too cryptic to be collected in quantity, or have a different habitat from larger crabs. Ovigerous females were present in all months except February (in both 1975 and 1976), but generally formed less than 50% of the total sample of females each month. The number of newly deposited eggs (0.75×0.81 mm) carried by females increased with increase in carapace width according to the equation y = -1123.56+102.97x (r ² = 0.8213). Egg development lasted 3–5 months, and egg mortality during this period was almost 10%. The overall sex ratio for the 18-month sampling period was 1846 ♀ : 993 ♂, which suggests that females were approximately twice as numerous as males. However, when crabs were sorted into size classes it was evident that the sex ratio was not significantly different from 1:1 in crabs of 6–12 mm carapace width, whereas females significantly (P<0.001) outnumbered males in the 13–21 mm size range. The right and left chela were approximately equal in length in females of all sizes, but the right chela of large males was greatly enlarged compared with the left, and with the chelae of females of comparable carapace width. Increase in the growth rate of the males’ right chela commenced at a carapace width of approximately 11.0 mm. Since the smallest ovigerous female collected also had a carapace width of 11.0 mm, it is concluded that both males and females attain sexual maturity at this size.     

Temporal changes in the reproductive population structures and males’ secondary sexual character of the hermit crab <Emphasis Type="Italic">Diogenes nitidimanus</Emphasis>

We investigated the reproductive ecology of D. nitidimanus in the Waka-River estuary with special reference to temporal change in the relative size of chelae length for males, i.e., secondary sexual character. Ovigerous females were observed from April to October, peaking in June–July with over 90% of females being ovigerous. Adult female carapace size ranged from 3.5 to 8.5 mm, but with the majority of females falling between 5–6 mm. Male carapace length was more evenly distributed between 3.5 and 10 mm. Juvenile settlement occurred mostly in July, during which time the frequency of both large females (over 6.5 mm in carapace length) and large males (over 8.5 mm in carapace length) clearly decreased. The carapace length of precopulatory-guarded females varied from 4.8 to 8.0 mm, while guarding males were almost over 7 mm and always larger than their paired females. The relative growth of the major chelae differed significantly between small and large males during the early months of the year, including the reproductive peak months (April–June). During these early months, large males had relatively larger chelae for their body size than did small crabs. This difference, however, was not evident later in the year (July–September). Large males may grow their chelae relatively long in the early months in order to take advantage of the mating opportunities during April–June. This is the first report in animals, to our knowledge, that relative size of the secondary sexual character for males temporarily change during a single reproductive season.     

Choose Your Weapon: Defensive Behavior Is Associated with Morphology and Performance in Scorpions

Morphology can be adaptive through its effect on performance of an organism. The effect of performance may, however, be modulated by behavior; an organism may choose a behavioral option that does not fully utilize its maximum performance. Behavior may therefore be decoupled from morphology and performance. To gain insight into the relationships between these levels of organization, we combined morphological data on defensive structures with measures of defensive performance, and their utilization in defensive behavior. Scorpion species show significant variation in the morphology and performance of their main defensive structures; their chelae (pincers) and the metasoma (“tail”) carrying the stinger. Our data show that size-corrected pinch force varies to almost two orders of magnitude among species, and is correlated with chela morphology. Chela and metasoma morphology are also correlated to the LD50 of the venom, corroborating the anecdotal rule that dangerously venomous scorpions can be recognized by their chelae and metasoma. Analyses of phylogenetic independent contrasts show that correlations between several aspects of chela and metasoma morphology, performance and behavior are present. These correlations suggest co-evolution of behavior with morphology and performance. Path analysis found a performance variable (pinch force) to partially mediate the relationship between morphology (chela aspect ratio) and behavior (defensive stinger usage). We also found a correlation between two aspects of morphology: pincer finger length correlates with the relative “thickness” (aspect ratio) of the metasoma. This suggests scorpions show a trade-off between their two main weapon complexes: the metasoma carrying the stinger, and the pedipalps carrying the chelae.     

Sex and Weapons: Contrasting Sexual Dimorphisms in Weaponry and Aggression in Snapping Shrimp

Sexual dimorphisms in weaponry and aggression are common in species in which one sex (usually males) competes for access to mates or resources necessary for reproduction – sexually dimorphic weaponry and aggression, in other words, are frequently the result of intrasexual selection. In snapping shrimp, the major chela (snapping claw) can be a deadly weapon, and males of many species have larger chelae than females, a pattern readily interpreted as resulting from intrasexual selection. Thus, males might be expected to show more sex‐specific aggression than females, and be more aggressive overall. We tested these predictions in two species of snapping shrimp in a territorial defense context. Neither of these predictions was supported: in both species, females, but not males, engaged in sex‐specific aggression and females were more aggressive than males overall. These contrasting sexual dimorphisms – larger weaponry in males but higher aggression in females – highlight the importance of considering the function of weaponry and aggression in contexts other than direct competitions over mates. In addition, species differences in the degree of sexual dimorphism in chela size were due to differences in female, not male, chela size, and the species with greater sexual dimorphism in weaponry was significantly less aggressive overall; also, while paired and solitary males did not differ in residual chela size, for the species with greater sexual dimorphism, females carrying embryos had smaller residual chela sizes. These results suggest that understanding the sexual dimorphisms in weaponry and aggression in snapping shrimp requires understanding the relative costs and benefits of both in females as well as males.     

Notes on the biology of Cancer bellianus (Brachyura, Cancridae) around the Canary Islands

Information on biometric and biological parameters of Cancer bellianus Johnson, 1861 (Crustacea, Decapoda, Brachyura, Cancridae) off the Canary Islands is given. Crabs examined were collected during experimental fishing surveys during 1974–1998. Carapace length, carapace width, total wet weight, sex and ovigerous condition were determined. This species was caught at depths from 153 to 750 m, the deepest ever recorded. Size frequency distributions were assembled and size-weight relationships were estimated by sex. Sex-ratio as a function of size and depth was determined. The size at first maturity was calculated by analysing the relative growth between the carapace length and the left chela width: 103.5 mm CL in males, 101.2 mm CL in females. Ovigerous females, egg size and fecundity estimates are reported apparently for the first time.     

Size,but not sex,predicts pinch force and exoskeleton mechanical properties in crayfish of the genus Faxonius

Studies of animal weaponry and defensive structures rarely take into consideration their underlying mechanical properties. We measured the compressive strength and thickness of the exoskeleton of the claw (chela) in two North American crayfish species, Faxonius virilis and F. limosus. We performed similar measures on the carapace, a body region not directly involved in agonistic contests. Males of both species generated significantly stronger maximum pinch forces than females. However, these differences can be attributed to differences in claw size between the sexes. The thickness (ultrastructure) of the claw exoskeleton was a significant predictor of its compressive strength and likely explained the difference in compressive strength we observed between the two species. Neither claw thickness nor claw compressive strength was correlated with maximum pinch force. Additionally, we found that crayfish body size was a strong predictor of carapace compressive strength and thickness, whereas sex was not. The claw had greater compressive strength and thickness than the corresponding values for the carapace. Our study shows that the mechanical properties of the crayfish exoskeleton are largely a function of size and highlights the need to integrate mechanical properties into studies of animal morphology and performance.     

Characterization of the exoskeleton of the Antarctic king crab Paralomis birsteini

Ocean acidification is projected to inhibit the biogenic production of calcium carbonate skeletons in marine organisms. Antarctic waters represent a natural environment in which to examine the long‐term effects of carbonate undersaturation on calcification in marine predators. King crabs (Decapoda: Anomura: Lithodidae), which currently inhabit the undersaturated environment of the continental slope off Antarctica, are potential invasives on the Antarctic shelf as oceanic temperatures rise. Here, we describe the chemical, physical, and mechanical properties of the exoskeleton of the deep‐water Antarctic lithodid Paralomis birsteini and compare our measurements with two decapod species from shallow water at lower latitudes, Callinectes sapidus (Brachyura: Portunidae) and Cancer borealis (Brachyura: Cancridae). In Paralomis birsteini, crabs deposit proportionally more calcium carbonate in their predatory chelae than their protective carapaces, compared with the other two crab species. When exoskeleton thickness and microhardness were compared between the chelae and carapace, the magnitude of the difference between these body regions was significantly greater in P. birsteini than in the other species tested. Hence, there appeared to be a greater disparity in P. birsteini in overall investment in calcium carbonate structures among regions of the exoskeleton. The imperatives of prey consumption and predator avoidance may be influencing the deposition of calcium to different parts of the exoskeleton in lithodids living in an environment undersaturated with respect to calcium carbonate.     

Deception with honest signals: signal residuals and signal function in snapping shrimp

Animals in competitive interactions often assess the competitiveability of opponents using signals. Signals used in competitiveinteractions are generally predicted to be honest, but opento low levels of deceit. Such "incomplete honesty" in signalscan be studied by using signal residuals, the residuals fromthe regression of a measure of signal structure on competitiveability. Specifically, individuals with positive signal residualsproduce signals that exaggerate their competitive ability; deceptive use of these signals may occur if signalers for whom the signalexaggerates their apparent competitive ability use the signalmore frequently. I used this framework to examine the use ofthe open chela display by big-clawed snapping shrimp (Alpheusheterochaelis). Competitive interactions between snapping shrimpare resolved primarily on the basis of body size, and the open chela display is used by males to assess body size. I foundthat the production of the open chela display by males respondingto superior competitors depends on chela residuals, such thatindividuals for whom the display exaggerates their apparentsize produce the display more often. This effect can be seenboth in response to isolated chelae and in staged competitiveinteractions. Interactions involving shrimp with larger chela residuals are long and highly escalated, suggesting that chelaresiduals affect assessment of competitive ability. Thus, theincreased use of the open chela display by males for whichthe display exaggerates apparent body size is an example ofdeceptive use of an otherwise honest signal.     

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.     

Cryptic asymmetry: unreliable signals mask asymmetric performance of crayfish weapons

Animals commonly use their limbs as signals and weapons during territorial aggression. Asymmetries of limb performance that do not relate to asymmetries of limb size (cryptic asymmetry) could substantially affect disputes, but this phenomenon has not been considered beyond primates. We investigated cryptic asymmetry in male crayfish (Cherax dispar), which commonly use unreliable signals of strength during aggression. Although the strength of a chela can vary by an order of magnitude for a given size, we found repeatable asymmetries of strength that were only weakly related to asymmetries of size. Size-adjusted strength of chelae and the asymmetry of strength between chelae were highly repeatable between environmental conditions, suggesting that asymmetries of strength stemmed from variation in capacity rather than motivation. Cryptic asymmetry adds another dimension of uncertainty during conflict between animals, which could influence the evolution of unreliable signals and morphological asymmetry.