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.     

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.     

Mechanisms of coexistence and competition between ants and land hermit crabs in a Bahamian archipelago

Ants and land crabs are common inhabitants of many coastal and insular communities across the tropics and subtropics, and yet direct evidence of interspecific competition between ants and land crabs has only recently been documented. I conducted a series of observational and manipulative experiments to further elucidate the mechanisms of competition, as well as coexistence, in these two groups in an archipelago of small Bahamian islands. Diel baiting trials demonstrated a significant temporal difference in foraging activity between the land hermit crab, Coenobita clypeatus (Herbst), and ant Brachymyrmex obscurior Forel, suggesting this is one mechanism underlying their coexistence on small oceanic islands. Reciprocal manipulative baiting experiments, in which one of a pair of species was removed from baits, documented that aggressive interspecific interactions underlie patterns of complementary distribution and temporal turnover at rich food resources. This was true for competition between hermit crabs and B. obscurior, and between B. obscurior and a second ant species, Dorymyrmex pyramicus Roger. Negative species associations at baits were found to be common throughout an archipelago of 69 small islands. A trade-off in exploitative and interference abilities may be a second mechanism allowing species coexistence on these small islands. Interspecific interactions such as competition and predation may occur commonly between ants and land crabs and have important consequences for the structure and function of tropical and subtropical insular ecosystems.     

Resource partitioning and competition for shells between intertidal hermit crabs on the outer coast of Washington

Summary Resource partitioning was quantified for 6 species of intertidal hermit crabs in the genus Pagurus, that occur on the outer coast of Washington. This, together with field evidence of shell shortage and with laboratory experiments to quantify the mechanism of interactions for shells, allowed estimation of the relative intensities of inter-and intraspecific competition between these species. The findings were that: (1) the magnitude of intraspecific competition was greater than any single interspecific competitive effect for all of the species; and (2) the relative proportion of intraspecific competition was greater for the middle and upper intertidal species than for the lower intertidal species. Studies at several outer coast sites supported these generalizations. Both of these findings are consistent with the hypothesis that competitive divergence has occurred in the past. The structure of the outer coast hermit crab assemblage is compared with that of the San Juan Archipelago hermit crab assemblage. Differences between the two do not seem to be the result of adaptive responses to the presence of more competing species in the former group.     

Frequencies of interspecific shell exchanges between hermit crabs

Frequencies of interspecific shell exchange due to shell fighting were determined for a number of species pairs of hermit crabs from several different locations. Frequencies were determined in the laboratory using a standardized experimental design. Results suggest that most individuals of most species are able to retain adequate or good quality shells in the presence of members of another species occupying poor quality shells. High frequencies of shell exchange always seem to be associated with very asymmetric relationships in which one member of the species pair is clearly dominant over the other. Dominant species usually attain larger sizes than subordinates, are found lower in the intertidal habitat, and are less abundant.     

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.     

Feeding hermit crabs to shrimp broodstock increases their risk of WSSV infection

White spot syndrome virus (WSSV) is a serious shrimp pathogen that has spread globally to all major shrimp farming areas, causing enormous economic losses. Here we investigate the role of hermit crabs in transmitting WSSV to Penaeus monodon brooders used in hatcheries in Vietnam. WSSV-free brooders became PCR-positive for WSSV within 2 to 14 d, and the source of infection was traced to hermit crabs being used as live feed. Challenging hermit crabs with WSSV confirmed their susceptibility to infection, but they remained tolerant to disease even at virus loads equivalent to those causing acute disease in shrimp. As PCR screening also suggests that WSSV infection occurs commonly in hermit crab populations in both Vietnam and Taiwan, their use as live feed for shrimp brooders is not recommended.     

Evolutionary palaeoecology of symbioses between bryozoans and hermit crabs

Modern hermit crabs form associations with many organisms which encrust, bore into, or cohabit the living chambers of gastropod shells occupied by the crabs. Among these hermit crab symbionts are bryozoan species which develop massive, commonly multilayered, colonies encrusting hermit crab shells. These colonies extend the living chamber of the crab through a characteristic process of helicospiral tubular growth originating from the shell aperture. The scant information available on the ecology of Recent bryozoan‐hermit crab symbioses is reviewed. Symbioses have been recorded from intertidal to upper slope environments, and from tropical to cold temperate zones. None of the hermit crab species are obligatory symbionts of bryozoans, and the majority of the modern bryozoan species involved are also not obligatory symbionts. Fossil examples always lack the hermit crabs, which have a poor fossilization potential; however, the distinctive tubular growth pattern and other features of the bryozoans enable recognition of ancient examples of the symbiosis. The earliest inferred associations between bryozoans and hermit crabs date from the Mid Jurassic, but associations remained uncommon until the Neogene. A remarkably wide taxonomic diversity of Recent and fossil bryozoans are known or inferred symbionts of hermit crabs. The broad evolutionary pattern of the association demonstrates multiple originations of the symbiosis by bryozoans belonging to at least 5 cyclostome and 12 cheilostome families. Only the Miocene‐Recent cheilostome family Hippoporidridae has an evolutionary history closely tied to symbiosis with hermit crabs. There is no evidence for coevolution.     

Neuromuscular correlates of rhythmical cheliped flexion behavior in hermit crabs

Journal of Comparative Physiology A -     

Effects of shell stress on the growth of hermit crabs

Growth rates of the hermit crabs Pagurus longicarpus Say and P. pollicaris Say maintained in preferred shells and in shells smaller than the preferred size have been compared. Changes in wet weight, shield length, and duration of the molt cycle were determined. Crabs in preferred shells grew significantly faster than those in small shells. P. pollicaris molted at approximately the same rate in both cases but grew more each molt in preferred shells. Von Bertalanffy growth curves were fitted to changes in shield length among crabs in preferred shells. These curves indicate that P. longicarpus may mature four months after settling from the plankton and reach its asymptotic size within the next eight months while P. pollicaris also matures four months after leaving the plankton but does not reach its asymptotic size for approximately three years. The rapid growth of P. longicarpus may enable it to preempt shells which are required for the successful brooding of a large clutch before these shells are required by more aggressive competitors, such as P. pollicaris and Clibanarius vittatus.     

Differential tolerance of body fluid dilution in two species of tropical hermit crabs: not due to osmotic/ionic regulation

The tropical intertidal hermit crabs Clibanarius taeniatus and Clibanarius virescens were examined for differences in survival and physiological responses in low salinity. We found that C. taeniatus survived better in dilute seawater than C. virescens and that these species did not differ in their abilities to regulate haemolymph osmolarity, ionic concentration of the haemolymph or body fluid volume. We also found no difference in oxygen consumption between the species when acutely exposed to a range of temperature and salinity combinations. It is concluded that the greater survival in dilute seawater by C. taeniatus compared to C. virescens is due to a greater tolerance of dilution of body fluids by C. taeniatus. Differences in tolerance to dilute seawater may influence the habitat preferences of these species within the same geographical area.     

Effect of gastropod shell characteristics and hermit crabs on shell Epifauna

The epifauna on gastropod shells occupied by the hermit crabs Pagurus pollicaris (Say) and P. longicarpus (Say) was examined, as was the utilization of shells by these two hermit crabs. In the study area in Tampa Bay, Florida, shells were not a limiting factor to the hermit crab population, and there apparently was little competition for shells. Interspecific competition for shells was limited because the two hermit crab species differed in size and hence occupied shells of different sizes. The total number and density of most epifaunal species were higher on shells occupied by hermit crabs than on unoccupied shells, possibly because hermit crabs prevent their shells from being buried and hence lengthen the time the epifaunal community can grow and develop. The hermit crab species also appeared to affect the epifaunal community, for the total number and density of most epifaunal species were larger on shells occupied by P. pollicaris than P. longicarpus. With increasing shell size, the populations of most epifaunal species, also were larger but not their density. Least influential in affecting the epifaunal community was the species of shells.     

Effects of chemical context on shell investigation behavior in hermit crabs

Specific chemicals in the environment evoke significant changes in the behavior of many aquatic organisms. We studied in the laboratory whether satiated individuals of the hermit crab, Pagurus longicarpus Say 1817, adjust their investigatory behavior towards an empty, optimal gastropod shell according to differences of chemical context. We also explored to what extent shell investigation by a crab in the same hunger state was affected by occupying an inadequately sized shell. Our results confirmed in part previous findings that crabs can discriminate the odor of freshly dead snails from the odor of freshly dead conspecifics. In the presence of the former odor, crabs inhabiting shells of inadequate size were more responsive and active than those in better-fitting shells. To the contrary, regardless of the quality of the inhabited shell, P. longicarpus remained practically motionless when presented with the odor of freshly dead conspecifics, possibly because the risks of incurring in predators would outweigh the benefits of acquiring a new shell. Unexpectedly, we found that crabs in both types of shell quality exhibited nearly the same behavior in control water, while crabs in adequate shells were more responsive in the presence of food odor. Individuals appeared insensitive to the odor of live snails; indeed, only one hermit crab species has been seen removing living snails from their shells. An intriguing result was that water conditioned by the odors of live conspecifics exerted a strong effect on all the individuals by inducing an intense shell investigation. Our study underlines the central role exerted by chemical detection in hermit crabs' behavior and demonstrates the existence of a complex interplay among chemical context, the physiological state of the animal, and the ecological pressures of the habitat.     

Resource assessment in hermit crabs: the worth of their own shell

Animals gather information about the quality of a resource throughits assessment and behave accordingly as a result of adaptivemotivational changes. In the hermit crab Pagurus longicarpus,we investigated whether an individual was affected in its motivationto acquire a new shell by the quality of the domicile shell(own resource value [ORV]), of the offered shell (external resourcevalue [ERV]), or of both and asked whether its motivation wasaltered by the information gathered during shell investigation.We analyzed the behavior of hermit crabs inhabiting shells ofdiffering qualities and compared their willingness to acquirean offered shell—optimal, smaller than optimal, or largerthan optimal—by measuring the latency to approach it,the number of shell investigation, and its total duration. Crabsin smaller shells (SSs) approached more quick and often theoffered shell, whereas crabs in larger shells investigated theoffered shell more thoroughly. The readiness of crabs to approachthe offered shell and the extent of its investigation were independentof the ERV but were exclusively affected by the ORV, whereasthe number and duration of shell investigation did not changewith time as investigation proceeded, except for crabs in SSs.These results suggest that P. longicarpus' motivation to acquirea new shell is exclusively influenced by the value of the shellit inhabits rather than by the quality of the shell it is offeredand that this species does not gather—or does not use—informationabout ERV during investigation.     

Multigene molecular systematics confirm species status of morphologically convergent Pagurus hermit crabs

Introduction

In spite of contemporary morphological taxonomy appraisals, apparent high morphological similarity raises uncertainty about the species status of certain Pagurus hermit crabs. This is exemplified between two European species, Pagurus excavatus (Herbst, 1791) and Pagurus alatus (Fabricius 1775), whose species status is still difficult to resolve using morphological criteria alone.

Methodology/Principal Findings

To address such ambiguities, we used combinations of Maximum Likelihood (ML) and Bayesian Inference (BI) methods to delineate species boundaries of P. alatus and P. excavatus and formulate an intermediate Pagurus phylogenetic hypothesis, based upon single and concatenated mitochondrial (cytochrome oxidase I [COI]) and nuclear (16S and 28s ribosomal RNA) gene partitions. The molecular data supported the species status of P. excavatus and P. alatus and also clearly resolved two divergent clades within hermit crabs from the Northeast Atlantic Ocean and the Mediterranean Sea.

Conclusions/Significance

Despite the abundance and prominent ecological role of hermit crabs, Pagurus, in North East Atlantic Ocean and Mediterranean Sea ecosystems, many important aspects of their taxonomy, biology, systematics and evolution remain poorly explored. The topologies presented here should be regarded as hypotheses that can be incorporated into the robust and integrated understanding of the systematic relationships within and between species of the genus Pagurus inhabiting the Northeast Atlantic Ocean and the Mediterranean Sea.