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.     

Shell fights in the hermit crabPagurus geminus: Computer simulation and experiment

The shell exchange in the hermit crabPagurus gemiuns was examined by computer simulations and experiments, to learn whether it is based on mutualistic interactions with both participants gaining, or on one-sided interactions with the large crab gaining regardless of loss and gain of the partner. The result of the experiment showed a much better fit to that of the one-sided simulation than the mutualistic simulation, suggesting that competitive interactions in which small crabs lose in the resulting shell exchanges are a part of shell fights of this species. While the shell exchange attempt can be regarded as aggressive, a higher probability of mutualistic encounters and of successful mutualistic shell exchanges in hermit crabs is discussed.     

Analysis of repeated signals during shell fights in the hermit crab Pagurus bernhardus

Shell exchanges between hermit crabs may occur after a period of shell rapping, when the initiating or attacking crab brings its shell rapidly and repeatedly into contact with the shell of the non-initiator or defender, in a series of bouts. There are two opposing models of hermit crab shell exchange and the function of shell rapping. The negotiation model views shell exchange as a mutualistic activity, in which the initiator supplies information about the quality of its shell via the fundamental frequency of the rapping sound. The aggression model views shell rapping as either detrimental to the defending crab, or as providing it with information about the initiator''s ability or motivation to continue, or both. The negotiation model makes no predictions about the temporal pattern of rapping, but under the aggression model it would be expected that crabs that rapped more vigorously would be more likely to effect an exchange. Repeating the signal could be expected under either model. Crabs that achieve an exchange rap more vigorously, rapping is more persistent when a clear gain in shell quality may be achieved, and the vigour is greater when the relative resource-holding potential (or ''fighting ability'') is high. These findings support the aggression model rather than the negotiation model. Contrary to the predictions of game theory, crabs that do not effect an exchange appear to signal that they are about to give up. The data suggest that rapping is performed repeatedly because the accumulation of all of the performances acts as a signal of stamina.     

Why small hermit crabs have large shells

Frequent shell exchanges among hermit crabs imply the enigmatic circumstance that large crabs frequently obtain large shells from smaller crabs. This seeming anomaly is explored as a key to the shell resource system. It is hypothesized to reflect how, where, and how often shells become available to the crabs. Shells become available infrequently, as snails die, and are available to the crabs for only a brief time before they become inaccessible. The standing crop of empty shells is almost always low and is irrelevant to rates of shell turnover in the crab population. Crabs are most likely to encounter shells of the wrong size, and the chance of encountering a shell of the desired size decreases as a crab grows. Snails and crabs are usually found on different portions of the shore; thus, crabs must make “foraging trips” for shells. Under this regime of shell supply, a crab will get a suitable shell the fastest when it accepts any fresh shell that is larger than its initial shell. It can then trade with other crabs to improve its shell fit. This behavior will make small crabs into a regular source of large shells for large crabs, and a shell exchange ritual will be strongly favored because both participants will benefit. Shells are an unusual resource because they are the object of both competitive and mutualistic interactions. This ambiguous quality is revealed in the intraspecific and interspecific responses of crabs to each other and to shells.     

Influence of prior experience on shell selection by the white spotwrist hermit crab Pagurus criniticornis (Crustacea: Paguridae)

Individuals of Pagurus criniticornis in a free-choice situation were experimentally tested under different laboratory conditions. In order to assess the effect of recently occupied shells on the size- and type-preference by hermit crabs, individuals were held for 30 days under one of the following two conditions: (1) excess of shells and (2) absence of shells. The crabs were then allowed to select shells from a wide array of empty gastropod shells of the two most-occupied species, as observed previously in the field: Cerithium atratum and Morula nodulosa. Preferred shell type (species) and size (shell aperture width and length) were correlated with hermit-crab size. The crabs showed a strong (100%) preference for C. atratum shells, demonstrating that recent and past experience did not influence either shell-type or shell-size preferences in this pagurid. Handling editor: K. Martens     

An analysis of competitive interactions between 3 hermit crab species

Summary Competition for empty gastropod shells in a group of three sympatric hermit crabs (Pagurus hirsutiusculus, Pagurus granosimanus, and Pagurus beringanus) was studied in the San Juan Archipelago, Washington State. Estimates of the competitive effects of each species on the others' shell supplies were derived using field data on shell utilization and the results of laboratory experiments to determine rates of acquisition and exchange of shells and preferences for different shell species. Each species experienced approximately an order of magnitude more intraspecific competition than interspecific competition for empty shells. This resulted from differences in preference for shell shapes, shell size use, and habitat use between P. hirsutiusculus and P. granosimanus, and largely from differences in habitat use between P. beringanus and the other two species. Experiments involving the release and recensusing of marked empty shells were used to estimate competitive effects more directly for the interaction between P. hirsutiusculus and P. granosimanus. Results were consistent with the estimates derived from data on resource partitioning. Possible causes of the low levels of interspecific competition are discussed, and results are compared with studies of other organisms that estimated both inter- and intra-specific competition.     

Individual distance in crustacea: IV. Distance and dominance hierarchies in Pagurus pollicaris

The agonistic acts and distances separating interacting individuals when acts occurred were recorded for specimens of the hermit crab Pagurus pollicaris. Individuals were observed in groups of six under several conditions (during a snail predation, with food in the aquarium, and nothing in the aquarium except the crabs before and after the crabs had access to empty gastropod shells).

The pattern of agonistic interactions within the groups showed a high degree of order, but the lack of changes in behavior over time indicated these crabs have an aggressive order rather than a dominance hierarchy.

The distances separating crabs when acts were executed were significantly affected by (1) the act being executed, (2) the act executed previously by the other crab, (3) the size of the acting crab, (4) the relative size difference of the two interactants, (5) the individual crabs’ identities, and (6) the condition of observation. The individual crab effects on distance were present when crab size was controlled and crabs that reacted at smaller distances tended significantly to elicit reactions at greater distances.     

Associations between gastropod shell characteristics and egg production in the hermit crab Pagurus longicarpus

Summary Two populations (salt marsh and seagrass bed) of the hermit crab Pagurus longicarpus were sampled to examine associations between shell characteristics and egg production. Multivariate statistical analyses controlled for crab size and time of year, variables that otherwise could be confounded with shell effects. Although correlations between shell characteristics and reproduction existed in both populations, generalizations could not be made because associations varied within and between populations. Shell species was not associated with a female's reproductive state (i.e., whether or not she was barren when sampled) in either population. In the seagrass population, medium-large and large females occupying severely damaged or fouled shells were half as likely to be reproductive as females occupying other shell conditions. However, there was no association between shell condition and reproductive state for small and small-medium females in the seagrass population or among all females in the salt marsh population. In the seagrass population, small through medium-large reproductive females occupied shells more similar to their predicted shell size, regardless of whether the occupied shell was relatively small or large, than nonreproductive females. In contrast, relative shell sizes of reproductive and nonreproductive females were similar for large females in the seagrass population and all females in the salt marsh population. Clutch sizes were enhanced for females occupying Polinices duplicatus shells or shells larger than their predicted shell size in the seagrass population. Relative shell size also was associated with clutch size in the salt marsh population, but crabs occupying shells similar to their predicted shell size had the largest clutches.     

Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers

Hermit crabs are critically dependent upon gastropod shells for their survival and reproductive fitness. While anecdotal reports have suggested that hermit crabs may be capable of removing live gastropods from their shells to access the essential shell resource, no systematic experiments have been conducted to investigate this possibility. This paper reports experiments on both marine (Pagurus bernhardus) and terrestrial (Coenobita compressus) hermit crabs in which crabs were paired in the laboratory with the gastropods whose shells they inhabit in the field. Pairings included both shelled and naked crabs and spanned the full range of the gastropod life cycle. Neither marine nor terrestrial hermit crabs were successful at removing live gastropods from their shells. Furthermore, only a small fraction of the crabs (5.7%) were capable of accessing shells in which the gastropod had been killed in advance, with its body left intact inside the shell. Finally, although hermit crabs readily entered empty shells positioned on the surface, few crabs (14.3%) were able to access empty shells that were buried just centimeters beneath them. These results suggest that hermit crabs are constrained consumers, with the shells they seek only being accessible during a narrow time window, which begins following natural gastropod death and bodily decomposition and which typically ends when the gastropod's remnant shell has been buried by tidal forces. Further experiments are needed on more species of hermit crabs as well as fine-grained measurements of (i) the mechanical force required to pull a gastropod body from its shell and (ii) the maximum corresponding force that can be generated by different hermit crab species' chelipeds.     

Quality of shells occupied by the hermit crabPagurus geminus: How many hermit crabs are satisfied with their shells?

The rate of changing shells in the hermit crabPagurus geminus was investigated to determine how many hermit crabs are satisfied with their shells. Animal collected from the coast of Oya, Tanabe, Wakayama Prefecture, were presented with fresh gastropod shells newly made by removing the soft parts. Approximately 80% of the hermit crabs changed from their original shells acquired in the natural habitat into fresh shells experimentally given to them and, thus, were regarded as dissatisfied with the shells possessed in the natural habitat. This condition was thought to lead to the fact that hermit crabs occasionally attempted to exchange shells between 2 individuals and even to attack living snails in the natural habitat.     

Same resource,different benefits: hermit crab shell structure advantages owners,but not intruders in agonistic interactions

Resources may confer advantages by enhancing their owners’ fighting ability (resource holding potential; RHP). While the resource-correlated RHP hypothesis has been recognized as a determinant of agonistic success in different taxa, this has mostly been based on assessment of either the intruder or the owner, but only rarely in both contestants. We tested whether the internal structure of shells affects hermit crabs’ RHP, both as owners defending the shell against eviction and as intruders attempting to gain access to an occupied shell. We conducted contests (n?=?60) to compare the success in shell eviction by intruders in intact shells vs. in shells with the columella artificially reduced, and the success of shell retention by owners in intact shells vs. shells with the columella reduced. The internal configuration of the shell showed different resource-correlated RHP effects depending on the individual’s role in the fight. The presence of a columella in the intruder’s shell did not affect the likelihood that they would evict their opponents. However, owners resisted more evictions in shells with intact columella than those in shells with reduced columella. Our results demonstrate that the same resource can offer different RHP advantages to owners and intruders during an agonistic interaction.

     

Antipredator defence of the hermit crab Pagurus filholi induced by predatory crabs

Hermit crabs have two antipredator tactics: taking refuge in its shell and fleeing. We examined the following two hypotheses using the hermit crab Pagurus filholi : (1) hermit crabs change their preference for shell types that they take refuge in and/or change the timing of fleeing (i.e. the duration of refuge in the shell) when they perceive a predator threat; (2) the type of shell that a hermit crab occupies affects the fleeing tactic of the individual. Under the stimulus of a crushed conspecific, hermit crabs changed neither their preference for shell species nor their refuge duration. On the other hand, under the stimulus of the predatory crab Gaetice depressus , hermit crabs increased their preference for Batillaria cumingi shells, which provide superior protection against predators, and shortened their refuge duration in the shells even when they occupied those effective against predation. Refuge duration was longer in B. cumingi shells than in the more vulnerable shells of Homalopoma sangarense . These results suggest that both antipredator defences (changing shell and timing of fleeing) are induced by the stimulus of a predator, and the timing of fleeing is affected by the shell type occupied.     

Interference and exploitation components in interespecific competition between sympatric intertidal hermit crabs

This study was designed to evaluate the effect of interference and exploitation competition in shell partitioning between two hermit crab species (Pagurus criniticornis and Clibanarius antillensis). Field samples revealed that shells of the gastropod Cerithium atratum were the main resource used by both hermit crab species and that Pagurus used eroded or damaged shells in higher frequency than Clibanarius. The exploitative ability of each species was compared between species in the laboratory using dead gastropod (Cerithium) baits to simulate predation events and signalize newly available shells to hermit crabs. Pagurus reached the baits more rapidly than Clibanarius, but this higher exploitative ability did not explain shell utilization patterns in nature. Another experiment evaluated the dominance hierarchy between these two hermit crab species and revealed that Clibanarius was able to outcompete Pagurus for higher quality shells in agonistic encounters. This higher interference competitive ability of Clibanarius in relation to Pagurus may explain field observations. Nevertheless, Pagurus may be responsible to enhance shell availability to other hermit crab species that have lower ability to find and use newly available shells. Differently, the poorer condition of shells used by Pagurus, the higher ability of this species to attend gastropod predation events and its higher consumption rate by shell-breaking crabs (Menippe nodifrons) may increase its predation risks, thus revealing the disadvantages of such an exploitative competitive strategy for hermit crabs.     

The power of shell rapping influences rates of eviction in hermit crabs

Hermit crabs fight for ownership of shells, and shell exchangemay occur after a period of shell rapping, involving the initiatingor attacking crab bringing its shell rapidly and repeatedlyinto contact with the shell of the noninitiator or defender,in a series of bouts. The temporal pattern of rapping containsinformation about the motivation and/or relative resource holdingpotential (RHP) of the initiator and acts as a repeated signalof stamina. Here we investigated the role of the force withwhich the rapping is performed and how this is related to thetemporal pattern of rapping by rubberizing the external surfaceof shells. Initiators that are prevented from rapping withtheir usual level of force persist with the activity for longer over the whole encounter but use fewer raps per bout and areless likely to effect an exchange than those supplied withcontrol shells. The fact that the force of rapping affectsthe likelihood of a crab being victorious suggests that eitherthe force of rapping contains information about motivation orRHP or that force directly affects noninitiators, reducingtheir ability to maintain an adequate grip on their shells.The data suggest that shell rapping is an agonistic signalrather than one that provides information useful to the noninitiator,as has been suggested by the negotiation model of shell exchange.     

Effects of shell status and social context on the agonistic behavior of the tropical hermit crab,Clibanarius signatus

In the hermit crabClibanarius signatus, the influence of both shell status and social context on agonistic behavior was examined. The experimental hermits were of the same size and sex, but differed in the status of the inhabited shell, which was either optimal, sub-optimal or damaged. The evidence from this experiment suggests that the agonistic behavior ofC. signatus is influenced to a great extent by asymmetries in the resource value; attacks are mainly performed by crabs which inhabit shells in a relatively poor condition and are mostly directed towards opponents with better-fitting shells. However, this effect differs according to the social context, being more pronounced when the potential gain is greater. Shell fights are decidedly more complex than most animal combats; more work is certainly needed to understand the mechanisms underlying resource assessment and decisions made during hermit crabs' interactions.     

Larval release behaviors of the striped hermit crab, Clibanarius vittatus (Bosc): Temporal pattern in hatching

Ovigerous hermit crabs, Clibanarius vittatus (Bosc), were examined in the laboratory to (1) determine if the time of larval release is a synchronous event, (2) determine the influence of a damaged gastropod shell during the egg hatching process, and (3) describe larval release behaviors. Ovigerous hermit crabs from natural light:dark (LD) and tidal cycles were moved to constant conditions 2-3 days prior to the predicted time of larval release. Larval release was synchronous, occurring near the time of expected sunset. Females with early-stage embryos placed under constant conditions displayed a free-running circadian rhythm, suggesting that the rhythm is under endogenous control. Hermit crabs with early-stage embryos that were placed under a shifted LD cycle (advanced 12 h relative to the ambient photoperiod) before being placed under constant conditions advanced their larval release rhythm by 12 h, indicating the rhythm can be entrained by the LD cycle. Hermit crabs with an intact shell released larvae in bursts at sunset over several consecutive nights (period = 24.2 h). In contrast, hermit crabs with damaged shells released larvae at different times over the course of a single day. Ovigerous females with intact shells exhibit several stereotypical hatching behaviors. The female stands on her walking legs and thrusts her abdomen, moving the shell in an oscillating motion. This movement may assist in breaking the outer membrane of the egg case. The female generates a water current inside the shell with her scaphognathite and mouthparts, which transports the newly hatched larvae out of the shell. Females in damaged shells did not display these behaviors; instead, larval release was a prolonged event with little movement of the female, and often the newly hatched larvae were not viable. These results indicate that an intact shell plays an important role in the hatching process for this hermit crab.     

Response of hermit crabs to sinistral shells

Shell rotating behavior of the hermit crabPagurus geminus was investigated. In preliminary observations, hermit crabs motivated to change shells rotated presented shells, filled with sand, in a way that dislodged the inside material. In order to determine if this behavior is stereotyped, or flexible and dependent on shell type, hermit crabs were tested with ordinary dextral shells ofLatirulus nagasakiensis and sinistral shells ofAntiplanes contraria. Sinistral shells are not normally encountered by hermit crabs. Their rotation of the dextral shell to the left was adequate for sand discharge. Sinistral shells were rotated in both directions. Analysis of recorded videotapes showed that variation in rotation direction could be attributed to variation in the position of the crab relative to the shell. When the crab faced the shell aperture from the inner lip, it rotated the sinistral shell to the right, and to opposite direction when it faced from the outer lip side. The crab always pushed the upper side of the horizontally laid shell, regardless of shell type or its own position.     

Shell recruitment in the Mediterranean hermit crab Clibanarius erythropus

Gastropod shells are vital for the majority of hermit crab species, being essential for their survival, growth, protection, and reproduction. Given their importance, shells are acquired and transferred between crabs through several modalities. We conducted observations and experiments at the Asinara Island (Sardinia, Italy) to investigate the efficacy of the different behavioral tactics adopted by the hermit crab Clibanarius erythropus to acquire shells, such as: (1) locomotion and activity at different tidal phases; (2) attendance at shell-supplying sites (simulated predation sites with five different odors: live and dead gastropods, live and dead crabs, predator); and (3) interactions with conspecifics in aggregations on simulated gastropod predation sites. In each tidal phase, locomotion was slow (0.7 cm min^− 1) and, as a consequence, the probability of encountering empty shells and conspecifics was low. Simulated gastropod predation sites quickly attracted a larger number of hermit crabs than the other sites tested. Aggregations seemed to function as shell exchange markets, as previously suggested for other species: the first attendant took the experimental shell and a chain of shell exchanges among conspecifics followed. Our results show that, in C. erythropus, aggregation is the most efficient tactic for the acquisition of new shells, whereas in other species, such as Pagurus longicarpus, it is associated with exploitation ability due to the intense locomotion. The interspecific plasticity in hermit crabs' behavior is confirmed.     

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.     

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.