Decision rules, energy metabolism and vigour of hermit-crab fights

Aggressive interactions between animals are often settled by the use of repeated signals that reduce the risk of injury from combat but are expected to be costly. The accumulation of lactic acid and the depletion of energy stores may constrain activity rates during and after fights and thus represent significant costs of signalling. We tested this by analysing the concentrations of lactate and glucose in the haemolymph of hermit crabs following agonistic interactions over the ownership of the gastropod shells that they inhabit. Attackers and defenders play distinct roles of sender and receiver that are fixed for the course of the encounter. Attackers perform bouts of 'shell rapping', which vary in vigour between attackers and during the course of the encounter, and are a key predictor of victory. In contrast to the agonistic behaviour of other species, we can quantify the vigour of fighting. We demonstrate, to our knowledge for the first time, an association between the vigour of aggressive activity and a proximate cost of signalling. We show that the lactate concentration in attackers increases with the amount of shell rapping, and that this appears to constrain the vigour of subsequent rapping. Furthermore, attackers, but not defenders, give up when the concentration of lactate is high. Glucose levels in attackers also increase with the amount of rapping they perform, but do not appear to influence their decision to give up. Defenders are more likely to lose when they have particularly low levels of glucose. We conclude that the two roles use different decision rules during these encounters.     

Use of energy reserves in fighting hermit crabs

When animals engage in fights they face a series of decisions, which are based on the value of the contested resource and either their relative or their absolute fighting ability. Certain correlates of fighting ability or 'resource holding potential' such as body size are fixed but physiological correlates are expected to vary during the encounter. We examine the role of energy reserves in determining fight outcomes and parameters during 'shell fighting' in hermit crabs. During these fights, the two contestants perform very different roles of attacker and defender. We show that the balance of the total energy pool, in the form of glucose and glycogen, determines the ability of defenders to resist eviction from their shells. Low glucose in evicted defenders is not caused by depletion of energy reserves, rather mobilization of glycogen appears to be the result of a strategic decision about whether to resist effectively, based on the perceived fighting ability of the attacker. Attackers, however, always initiate the fight so such a decision for this role appears unlikely. In addition to influencing decisions and ability during fights, physiological correlates of fighting ability can in turn be influenced by strategic decisions.     

Shell fights in the hermit crabPagurus geminus: Effect of cheliped use and shell rapping

In shell fights of the hermit crab,Pagurus geminus, frequently it is observed that large crabs (attackers) grasp the thoracic appendage of small crabs (defenders) with the major cheliped and pull the smaller crabs out of their shells. If this is a standard occurrence and result, then the interaction should not be called a “negotiation” (Hazlett 1978). The role of cheliped use by the attckers in the eviction of defenders was therefore studied using crabs with tubes on their chelipeds, and the effect of shell rapping, which is thought to be necessary for eviction, was studied using crabs without shells. The experimental crabs evicted the defenders but fighting was significantly prolonged. Therefore, the negotiation model cannot be rejected. Specific aspects of shell fights in hermit crabs are discussed.     

Cumulative or sequential assessment during hermit crab shell fights: effects of oxygen on decision rules

Agonistic interactions between animals are often settled by the use of repeated signals which advertise the resource-holding potential of the sender. According to the sequential assessment game this repetition increases the accuracy with which receivers may assess the signal, but under the cumulative assessment model the repeated performances accumulate to give a signal of stamina. These models may be distinguished by the temporal pattern of signalling they predict and by the decision rules used by the contestants. Hermit crabs engage in shell fights over possession of the gastropod shells that they inhabit. During these interactions the two roles of signaller and receiver may be examined separately because they are fixed for the duration of the encounter. Attackers rap their shell against that of the defender in a series of bouts whereas defenders remain tightly withdrawn into their shells for the duration of the contest. At the end of a fight the attacker may evict the defender from its shell or decide to give up without first effecting an eviction; the decision for defenders is either to maintain a grip on its shell or to release the shell and allow itself to be evicted. We manipulated fatigue levels separately for attackers and defenders, by varying the oxygen concentration of the water that they are held in prior to fighting, and examined the effects that this has on the likelihood of each decision and on the temporal pattern of rapping. We show that the vigour of rapping and the likelihood of eviction are reduced when the attacker is subjected to low oxygen but that this treatment has no effect on rates of eviction when applied to defenders. We conclude that defenders compare the vigour of rapping with an absolute threshold rather than with a relative threshold when making their decision. The data are compatible with the cumulative assessment model and with the idea that shell rapping signals the stamina of attackers, but do not fit the predictions of the sequential assessment game.     

How resource quality differentially affects motivation and ability to fight in hermit crabs

Contesting animals typically gather information about the resource value and that information affects fight motivation. However, it is possible that particular resource characteristics alter the ability to fight independently of the motivation. Using hermit crabs, we investigate how the resource in terms of shell quality affects both motivation and ability to fight. These crabs fight for shells, but those shells have to be carried and may impose physiological costs that impede fight vigour. We find that the shell has different effects on motivation and ability. Potential attackers in very small shells were highly motivated to attack but, rather than having enhanced ability, unexpectedly quickly fatigued and subsequently were not more successful in the fights than were crabs in larger shells. We also examined whether defending crabs could gather information about the attacker''s shell from the vigour of the attack. Defending crabs gave up quickly when a potential gain had been assessed, indicating that such information had been gathered. However, there was no indication that this could be owing to the activity of the attacker and the information is probably gathered via visual assessment of the shell.     

Sexual dominance in hermit crab shell fights: asymmetries in owner–intruder status,crab size,and resource value between sexes

We studied sexual dominance and seasonal differences in aggressiveness of individuals in intraspecific competition for shells of the hermit crab Pagurus filholi in terms of size of contestants and duration of the attempt to deprive other crabs of their shell. Experiments were conducted using paired intrasexual and intersexual contests in the pre-breeding and post-breeding seasons. Size ratios between contestants were systematically varied to assess the sexual difference in size and owner advantages. In both intrasexual and intersexual contests intruder crabs tended to win the contests more often as their size increased, that is, size advantage overcame owner advantage. Although we did not recognize a sexual difference in size and owner advantages in contest outcomes, male intruder crabs took a shorter time to deprive female owners of a shell than to deprive male owners. Furthermore, male individuals in the pre-breeding season had significantly longer fight durations. Fighting is costly. Thus males can afford to expend more energy and time fighting, indicating that males are dominant over females in shell fights as both intruders and owners. Electronic Publication     

The Napoleon Complex: why smaller males pick fights

Does it ever pay for smaller animals to initiate fights even when they are likely to lose? Asymmetry in payoffs between opponents or a suboptimal strategy resulting from likely losers misperceiving themselves as likely winners have both been proposed as possible explanations for the aggressive behavior of smaller males. The model presented here suggests that in some cases, even without a payoff asymmetry and allowing for only a small error in perception, likely losers are expected to attack first. If the value of the resource exceeds the cost of losing a fight, the cost of displaying is sufficiently small, and assessment of resource holding power is reasonably accurate but not perfect, the evolutionarily stable strategy (ESS) prompts those contestants who perceive themselves as the likely losers to initiate fights, while it prompts those contestants who perceive themselves as the likely winners to wait for the adversary to attack or retreat.     

Shell fights in the hermit crabPagurus geminus: Aggressive act with mutual gain

The shell exchange of the hermit crabPagurus geminus was investigated to see whether it is based on negotiation (Hazlett 1978) or aggression (Elwood & Glass 1981) between the 2 interactants. Two conditions were adopted: the mutualistic condition in which both participants gain from resulting shell exchange, and the competitive condition in which one interactant gains at the expense of the other. The assumption is that if the interactants negotiate, then a higher, and/or quicker, rate of shell exchanges occurs in the mutualistic condition than in the competitive condition. Almost all the defenders (smaller crabs) were evicted from their shells in both of the conditions. No significant difference was detected in the fight time between the conditions, suggesting a low probability of negotiation. In the second experiment, the attackers (larger crabs) were deprived of the use of their chelipeds, allowing me to observe defenders' behavior when they were not subjected to forced eviction. In this case, the defenders evacuated their shells significantly more quickly under competitive conditions, but remained in the shell much longer under the mutualistic condition, indicating that the shell exchange of this species involves no negotiation. The function of shell rapping and the possibilitics of mutual gain in the aggressive interaction are discussed.     

Shell wars II: the influence of relative size on decisions made during hermit crab shell fights

The value of contested resources (shells) in hermit crab fights depends on the sizes of the crabs relative to the sizes of the resources. Thus when relative contestant size is the main experimental variable, motivational factors associated with shell size will also be an experimental problem. Two experiments are described that together overcome this problem. Relative crab size influences all stages of shell fights including pre-fight display, escalation, eviction and examination of the opponent's shell by the victor both before and after eviction of the loser. Shell fights occur more often between disparately-sized animals than between those similar in size. This apparent contradiction of recent theory (Maynard-Smith & Parker 1976) is probably due to the high cost of being without a shell and the small chance that an escalated fight will result. Relative crab size influences the time taken in resource assessments and thus the effectiveness of these assessments is also probably influenced. Causal factors influencing each of the major decisions in shell fights are described and although these fights are more complex than most they are in general agreement with theory on animal contests.     

The Role of Circulating Metal Ions During Shell Fights in the Hermit Crab Pagurus bernhardus

Fighting animals must make a series of decisions, and understanding the proximate causes behind these decisions can give insight into how they are made. For example, fights have been analysed with respect to energetic costs and endocrine changes associated with engaging. However, another mechanism for the control of demanding activity, such as fighting, is the modulation of aerobic capacity by divalent metal ions. Here we examine post‐fight haemolymph concentrations of magnesium (Mg²⁺) and calcium (Ca²⁺) ions in the common European hermit crab Pagurus bernhardus. Hermit crabs fight over the ownership of gastropod shells, where they adopt two very different roles during the encounter: attacker and defender. Despite the two roles performing different activities, we found that Mg²⁺ and Ca²⁺ affected them similarly, with concentrations of these ions being highest in successful individuals. Haemolymph concentrations of Mg²⁺ and Ca²⁺ were also found to increase as a result of fighting, and these elevated levels will, via allosteric interactions, increase the oxygen affinity of the respiratory pigment haemocyanin, enhancing respiratory capacity and therefore fighting ability. Furthermore, the present study revealed that seasonal changes in circulating levels, along with the ability of competitors to respond to them, may ultimately influence an individual’s success in aggressive interactions.     

Negotiation or aggression during shell fights of the hermit crab Pagurus bernhardus?

Shell fighting behaviour of the hermit crab Pagurus bernhardus was investigated. Analysis of fights between crabs in which there was little or no asymmetry in potential benefit for the two crabs from a shell exchange suggested that the duration of the fight increased as the potential benefit increased. Further experiments indicated that a naked crab was capable of evicting a housed crab by a process of direct aggression. Analysis of fights in which there was a slight asymmetry in potential gains from shell exchange indicated that the result of the fight was primarily determined by the large of the two crabs. These results are contrary to the proposal of Hazlett (1978) that the interactions represent a process of negotiation rather than aggression.     

Estimating the energetic cost of fighting in shore crabs by noninvasive monitoring of heartbeat rate

After establishing shore crabs, Carcinus maenas, individually in separate aquaria, we used a noninvasive infrared phototransducer to monitor their heartbeat rate continously before, during and after fights with intruder crabs. We confirmed that heartbeat rate is a reliable indicator of oxygen consumption and then used it to estimate indirectly the energetic cost of fights differing in duration and intensity, and its dependence on prior residence and relative size of opponent. Prior residence in aquaria significantly increased the probability that crabs would initiate fights against intruders. The majority of fights were resolved by aggressive contacts, display being used extensively only against smaller intruders. Fights between evenly sized opponents and between residents and larger intruders involved almost continuous aggression, whereas fights with smaller intruders involved several shorter bouts of aggression. Fight duration was weakly correlated with the relative size of opponents. Heartbeat rate, measured only in residents, was elevated above resting levels throughout fights, hence energy expenditure during fighting increased linearly with fight duration. Contrary to expectation, heartbeat rate was not significantly influenced by relative size of the opponent or by the intensity of aggression. After fighting, heartbeat rate usually returned to resting levels within 30-60 min, recovery taking longer in fights against larger intruders, when the fight was always lost. We propose that prolonged elevation of heartbeat rate in residents that had lost to larger intruders represented a state of alertness, adaptive against impending risks of resource loss or injury. Copyright 2000 The Association for the Study of Animal Behaviour.     

Imperfect assessment and limited information preclude optimal strategies in male-male fights in the orb-weaving spider Metellina mengei

Agonistic behaviour between male orb-web spiders Metellina mengei competing for access to female webs was examined in field experiments to test the major predictions of game theory. Winners of fights were significantly larger than losers, particularly with respect to the length of the first pair of legs, which are sexually dimorphic in this species and used extensively in agonistic encounters. The size of the winning male had no influence on contest intensity or duration, and neither did relative size. However, fight intensity and duration were both positively correlated with the size of the losing male. Resident males won significantly more contests than intruders. Winning intruders were significantly larger than winning residents and it was these winning intruders that tended to produce the longer fights. Female weight and hence reproductive value had a marked influence on fight intensity and duration of fights won by the intruder but not those won by the resident. This indicates that only the resident obtains information about the female. These data are discussed with reference to the discrepancy with theory and a failure of some contestants to obtain information on resource value and relative contestant size necessary to optimize fight strategy.     

Post‐Contest Stridulation Used Exclusively as a Victory Display in Mangrove Crabs

Victory or triumph display is a post‐contest signal, performed only by winners and not by losers. While much is unknown about its function, there is mounting evidence that victory displays are widespread among animals. However, evidence remains anecdotal in crabs. Sesarmid crabs belonging to the genera Parasesarma and Perisesarma are known to have characteristic stridulatory structures on their chelipeds. In Perisesarma eumolpe, a mangrove crab, stridulation has been anecdotally purported as a triumph display. We examined whether stridulation in P. eumolpe is a victory display and the factors affecting it by staging 17 contest trials among males and investigating the factors influencing stridulations and fight outcome in 55 fights. Using generalised linear mixed‐effects models, we find that stridulations were generally performed by winners and after fights, especially when the fights were intense. In addition, stridulation was only observed in the context of a contest, never before or outside of it. Stridulation in P. eumolpe is likely a victory display, and, unlike other forms of victory display described for other species, it appears exclusively used for asserting victory.     

Effect of melatonin on hemolymph glucose and lactate levels in the fiddler crab Uca pugilator

Melatonin was injected into intact and eyestalk-ablated fiddler crabs (Uca pugilator), and its effects on hemolymph glucose and lactate levels were studied. In intact crabs, glucose and lactate levels cycled simultaneously, with peaks occurring during early and late photophase. Melatonin caused a shift in the glucose and lactate cycles, with only one peak occurring closer to mid-photophase. In eyestalk-ablated animals, the glucose rhythmicity was lost; lactate cycled, but levels were significantly lower than in intact animals. Melatonin caused a delayed hyperglycemia in eyestalk-ablated animals, with concurrent but much lower increases in lactate. Overall, melatonin demonstrated delayed hyperglycemic effects that do not appear to be mediated solely via eyestalk factors such as crustacean hyperglycemic hormone (CHH), though involvement of the eyestalks cannot be ruled out. An influence on extra-eyestalk CHH secretion is a potential mechanism of melatonin activity.     

Functions of fights in territory establishment

Fights are often observed when prospective territory owners settle in patches of vacant habitat, but the function of these fights in space acquisition is obscure. This study tests two hypotheses about the effect of fights on subsequent space use patterns: first, that settlers win space by winning fights and, second, that fights encourage the establishment of mutually exclusive home ranges between opponents (i.e., "fights make neighbors"). The behavior of juvenile Anolts aeneus lizards was recorded as they established territories in patches of habitat in the field. In support of the fights-make-neighbors hypothesis, opponents whose last aggressive interaction was a fight were six times more likely to have mutually exclusive home ranges at the end of the settlement period than were otherwise equivalent dyads whose last encounter was a chase. Contra the hypothesis that settlers win space by winning fights, most last fights ended in a draw, and there was no discernable relationship between the outcome of last fights and the subsequent space use of the contestants. These and previous analyses of settlement behavior in this species suggest that fights during the settlement period encourage the formation of symmetrical social and spatial relationships between neighboring settlers.     

Initiation and Resolution of Fights between Swimming Crabs (Liocarcinus depurator)

Fights between male swimming crabs (Liocarcinus depurator) were studied in the laboratory and in the field. These crabs fight readily in the laboratory, interactions being initiated equally often by the larger and the smaller of the two opponents, but usually being won by the larger. Many different combinations of cheliped and swimming leg postures are seen during fights, which fell into 4 categories: 13% involved only stationary display, 38% were resolved by a single approach and retreat by displaying crabs, 9% were resolved by multiple approaches and retreats and 39% involved physical contact between the crabs. Fight length was variable, and depended both on the absolute size and on the relative size of the participants. During the course of the fights, few behavioural differences were observed between the eventual winners and losers. Fights were less common in the field, but the same rules determined their initiation, content and outcome. The results are discussed in the context of game theory.     

Is Difference in Body Weight,Antler Length,Age or Dominance Rank related to the Number of Fights between Fallow Deer (Dama dama)?

Competitive interactions between mature male fallow deer were investigated to determine whether antler length, body weight, age or dominance rank were related to the number of fights between individuals. Four different hypotheses were tested; the first predicted that body weight and antler length indicate individual quality and, therefore, as the difference between competitors in body weight and antler length increases, there should be a corresponding decrease in fight rate. The second and third hypotheses predicted that, as difference in dominance rank increased there would be a decrease in fight rate between males either, as a risk reduction measure or because of inhibitory control by dominant males. A fourth hypothesis predicted that, where dominance rank does not mediate fight rate, that similarity between contestants based on age might be important. If this is the case, then as the difference in age between competitors increased there should be a decrease in fight rate. Our results show that when dominance rank is controlled for, there was no relation between body weight, antler length or age with fight rate. There was a negative relation between dominance rank difference and fight rate, a result that supports both the risk reduction and inhibitory hypotheses. There was an increasing tendency to fight with closely ranked males as cohorts reached peak reproductive age; males aged 5 yr fought with other 5‐yr‐old males based on rank difference and males aged 6 yr fought with other males aged six and across all age groups based on rank difference. This trend was not observed in 4 or 7‐yr‐old males. Our results suggest that males in prime breeding condition limit potential costs of fighting (such as time and energy) by only interacting with other males of similar rank.     

Do weaponless males of the hermit crab <Emphasis Type="Italic">Pagurus minutus</Emphasis> give up contests without escalation? Behavior of intruders that lack their major cheliped in male–male contests

In dyadic contests, theoretical studies have predicted that weaker contestants are less likely to engage in fights to minimize the cost of aggression. Since the major cheliped of decapod crustaceans is critically important as a weapon, contestants without a major cheliped should be more likely to give up the contests. We therefore examined whether loss of the major cheliped by the hermit crab Pagurus minutus would affect their decision to escalate male–male contests over guarded females. Intruders without a major cheliped showed no difference in the frequency of escalation compared with intact intruders, and the decision to give up was affected by the body size difference between the contestants. After escalation, compared with intact intruders, intruders without a major cheliped had significantly decreased success of takeover of a female from opponents, suggesting a strong disadvantage of losing their major cheliped. Although the decision of weaponless intruders to escalate seems irrational, several factors, such as poor accuracy of resource holding potential assessment, the influence of body size, and a high benefit to cost ratio of male–male contests, may have affected their behavior.     

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

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