Shell selection in the hermit crab Pagurus bernhardus

Three shell variables were deemed to be important in the selection of a shell by the hermit crab Pagurus bernhardus (L.). These were the weight, volume and aperture width of a shell. Each of these variables were used in multiple regressions against crab weight for 25 Littorina littorea and 25 Thais lapillus shells chosen when both shell species were together in a tank, and for a further 25 L. littorea and 25 T. lapillus when the two shell species were separate. Principal component analysis was performed on the four groups of 25 shells selected, and multiple-regression equations were calculated using the principal components as the new variables. It was found that crabs chose a shell of suitable general dimensions rather than solely on the basis of one shell variable. The advantages of using statistical techniques developed in this paper over previous approaches to shell selection are discussed.     

Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Predaceous gastropods regulate new-shell supply to salt marsh hermit crabs

Marsh hermit crabsPagurus longicarpus Say directly acquire new shells as the predatory gastropodMelongena corona Gmelin consumes marsh periwinkles,Littorina irrorata Say. The influx rate of new shells into a salt marsh hermit crab population was measured by marking live periwinkles and daily recovering the shells from hermit crabs over periods of 3 to 6 d. Average rates of new shell acquisition ranged from 4.0 to 23.3 new shells per day from salt marsh areas of 4×10 m. Such consistently high rates contrast with the negligible rates generally assumed for new shell entry into hermit crab populations. The number of new shells acquired each day varied directly with the number of the predatory gastropod,M. corona, present in each study area at both natural and manipulated predator densities. Empty shells on the substrate are usually considered as the primary source of new shells to hermit crabs. However, over 500 empty shells had to be placed on the substrate in a 4×10 m area to provide a daily rate of 20 new shells to the hermit crab population.This is the first in a new contribution series from the Florida State University Marine Laboratory No. 1001     

Utilization of partially predated snail shells by the hermit crab Pagurus longicarpus Say, 1817

The hermit crab Pagurus longicarpus was shown to inhabit shells that were partially predated from intertidal areas of Long Island, New York. Among field collections of P. longicarpus, 2.13% of the hermit crabs (46 of 2155) were found with shells with snail tissue present. Over 90% of these partially predated snail shells were occupied by male hermit crabs. Although hermit crabs were in 8 species of snail shells, only Littorina littorea and Nassarius obsoletus were found occupied by hermit crabs and containing snail tissue. In the laboratory, we found that specimens of the spider crab Libinia emarginata were able to pull off the operculum of snails, leaving damage as found in field collections. In contrast, specimens of P. longicarpus were not able to prey on live, healthy snails. When specimens of P. longicarpus were placed in communal tanks, hermit crabs preferred partially predated snail shells to empty and original shells. However, original shells and empty shells were occupied with more frequency than partially predated shells when crabs were isolated. These findings indicate P. longicarpus actively seeks shells soon after attack and abandonment by snail predators, especially in the presence of competitors.     

Obligate commensalism of <Emphasis Type="Italic">Curvemysella paula</Emphasis> (Bivalvia: Galeommatidae) with hermit crabs

Curvemysella paula is a markedly crescent-shaped bivalve that lives inside snail shells occupied by hermit crabs. Here, we describe the unique symbiotic life, growth pattern, and reproductive biology of this bivalve, based on specimens collected from the shallow, muddy bottom of the Seto Inland Sea, Japan. C. paula was found attached to columellae in the siphonal canal, mainly of nassariid snail shells occupied by two types of hermit crabs: Diogenes edwardsii (Diogenidae) and Spiropagurus spiriger (Paguridae). The crescent-shaped shell of C. paula is an adaptation to symbiotic life in the narrow interspace between the snail shell and the hermit-crab abdomen. C. paula is a protandric hermaphrodite. In our samples, each host snail shell harbored one (or rarely a few) large female and several males. All the female bivalves settled on the host shells with their anterior end facing outward and benefited from currents created by the hermit crab when feeding. In the muddy bottom, snail shells are a limited resource for both the hermit crabs and symbiotic bivalves. The bivalves benefit from the mobility of the hermit crabs, which prevent the shells from becoming buried in the mud. C. paula represents the only example of obligate commensalism with hermit crabs found in Bivalvia.     

Epibiosis on cerith shells in a seagrass bed: correlation of shell occupant with epizoite distribution and abundance

The cerith Cerithium atratum (Born 1778) is an abundant gastropod in the seagrass beds at Cabo Frio, Brazil. In order to estimate the ecological importance of cerith shells as a rare hard substratum in the seagrass bed, the abundances of C. atratum and of cerith shells occupied by hermit crabs were quantified. The mean densities of C. atratum and hermit crabs were 1887 and 100 individuals m⁻², respectively, and these provided 0.5 m² shell area m⁻² available for epizoite colonization. The tube-forming polychaete Hydroides plateni (Kinberg 1867) and oyster Ostrea puelchana Orbigny, 1841 were the dominant visible epizoites on inhabited cerith shells. These epizoite populations were compared in order to investigate whether the temporal and spatial patterns in the epibiotic community were related to ecological and behavioral aspects of the occupant species (cerith or hermit crab). Larger cerith shells had a greater abundance of epizoites. Each epizoite showed a preference for a different occupant of the shells (the oyster for C. atratum and the polychaete for cerith shells occupied by hermit crabs). The oyster showed a seasonal pattern in abundance on C. atratum, being more common in fall (March–April). The distribution of the epizoites on the shells depended on the shell occupant species and was probably related to their different foraging activity –C. atratum ploughs half buried through the sediment, while the hermit crab crawls on the sediment surface. In both cases, the activity of the shell occupant was considered to be beneficial to the epizoites, as empty shells and shell fragments did not support a macroepifauna. Received: 1 May 2000 / Accepted: 8 August 2000     

Shell acquisition by hermit crabs: which tactic is more efficient?

Gastropod shells are limited resources for most hermit crab species, acting as primary factors in various aspects of their biology. To investigate the efficacy of different behavioral tactics adopted for their acquisition (locomotion, attendance at shell-supplying sites, interactions with conspecifics in aggregation) we conducted observations and experiments at a salt marsh in New England (USA). Locomotion, fast and meandering, significantly increased the chances of encountering empty shells and conspecifics. However, shell exchanges were rare. Simulated gastropod predation sites quickly attracted a larger number of hermit crabs than the other shell-supplying sites tested (live and dead conspecifics, live snails) and induced the rapid occupancy of all the empty shells offered, usually by the first crabs arriving at the site. Aggregations seemed not to function as shell exchange markets, as previously suggested for several other species. In the short run, exploitation seems to be more efficient for the acquisition of new shells by Pagurus longicarpus. In the long run, it is the density of nondestructive gastropod predators that regulates the availability of new shells of good quality in the pool available to this hermit crab species.     

Fighting behavior in hermit crabs: the combined effect of resource-holding potential and resource value in Pagurus longicarpus

A study was conducted to determine whether asymmetries in both resource-holding potential (RHP) and resource value (RV) influence dominance and fighting behavior in the hermit crab Pagurus longicarpus. A total of 120 groups of three crabs were observed for 10 min in four experiments that tested all diverse combinations of equal/different RHP (i.e. ‘body size’) and equal/different RV (i.e. ‘shell size’ and ‘shell quality’). In a fifth experiment, dominant and subordinate individuals of the same size category (26 groups) were forced to enter shells of opposite quality than those previously occupied, and then the behavior of the reconstituted original groups was observed for additional 10 min. As expected, crabs in lower quality shells were more willing to initiate and to escalate fights. However, their attacks were directed to any crab of the group, independently of the defender's shell quality, and the fight duration did not vary with the different value of the resources at stake. This may indicate that P. longicarpus is unable to assess the quality of the shells available in its social environment but bases its tactical decisions during fights solely on its own resource. This suggestion was confirmed by the change in the fighting behavior of crabs whose shell quality was experimentally altered. This manipulation induced an overall increase in the intensity of aggression, drastic modification of crab behavior, and inversion of the hierarchy even though these crabs have had previous experiences of wins/losses and were familiar to the other members of the group. In this species, large crab size and/or the occupancy of adequate (and oversized) shells appeared to be the most likely determinant of contest resolution. Individuals seemed to retain a memory of the previously held resource and behaved accordingly.     

Ecology of subtropical hermit crabs in SW Madagascar: short-range migrations

Many mobile animals migrate because of the different benefits provided by different localities in time and space. For hermit crabs, such benefits include resource (shell, water, food) acquisition and gamete release. One of the more successful crustacean land-invaders, Coenobita hermit crabs, undertake complex short-range migrations in SW Madagascar. Number of active hermit crabs was inversely related to wind strength and positively related to tidal range, emphasising that movement would conserve water. A circadian component was also recorded in the locomotory activity of Coenobita pseudorugosus and C. rugosus. Path linearity varied with many of the same parameters, but also with beach slope. Movement was primarily perpendicular to shore in small individuals, but the parallel proportion increased with hermit crab size and tidal range, probably driven mostly by shell and food searching. Despite the costs of movement and shell carriage in the terrestrial environment, C. pseudorugosus and C. rugosus were as fast as their marine counterparts. Their speeds varied principally with individual size and were approximately 20% faster without shells and about 20% slower when climbing up a 20° slope, compared to horizontal or downhill travel. Hermit crabs, which are highly numerous and speciose in SW Madagascar, do not seem to partition niches by differential movement patterns. Aside from provision of shells in middens and capturing large adults for bait or pets, human activity may have a profound effect on hermit crab movement: observations at rare uninhabited marine reserves like Nosy Ve show that considerable diurnal activity may take place despite the apparent hostility of the environment to an essentially marine animal.     

Body and resource size at the land–sea interface

Body size in animals varies with many parameters, amongst them taxonomic affiliation, lifestyle and ambient environment oxygen levels. Size has considerable implication to possibilities for animals; for example, parasites need to be small and top predators large. Body size and resource requirements (shell size) were investigated across the land–sea interface in hermit crabs (Crustacea: Malacostraca: Decapoda) and snails (Mollusca: Gastropoda: Prosobranchia). These are two of the few taxa to occur in the sea, on the shore and on land as residents. Both taxa are also appropriate for such an analysis as they are abundant, speciose, cohabit the same environments and are linked—gastropod shells are a critical resource to hermit crabs. Both the maximum and mean sizes of hermit crab species showed parabolic relationships with shore height, decreasing from the sublittoral and supralittoral to the eulittoral. Average maximum size of gastropods exhibited a similar intertidal minimum although variability was high. It is suggested that this pattern is robust: not only did two distantly related taxa show the same pattern, but neither region nor site contributed significantly to total variability. The mass of resources (gastropod shells) used by hermit crabs, however, showed a converse pattern. The smallest shells (relative to hermit crab body size) were used in the sublittoral and supralittoral. Response to environmental stress and predation pressure are offered as two alternate theories to explain the observed body dwarfism and resource gigantism in the intertidal zone.Communicated by J.P. Thorpe, Port Erin     

Consequences of shell-species preferences for female reproductive success in the hermit crab Pagurus bernhardus

Littoral hermit crabs, Pagurus bernhardus, show a strong preference for Littorina obtusata shells rather than those of Gibbula species. The fitness consequences, in terms of fecundity, for this shell preference is examined for female crabs. Females in the preferred species produced eggs earlier in the season, produced more eggs in the first brood, and produced a second brood more often than did females in the less preferred species. The smaller brood for Gibbula spp. was not a consequence of egg loss from the pleopods due to an unfavourable shape of shell, but rather reflected lower egg production. It is not clear, however, if this differential reproduction is due to direct costs of carrying an unfavourable shell, i.e. the shell impedes reproduction, or whether crabs compete aggressively for favoured shells so that only crabs of low quality inhabit lowquality shells.     

Shell preference in a hermit crab: comparison between a matrix of paired comparisons and a multiple-alternative experiment

The preference of the hermit crab, Calcinus californiensis, among six species of shells, was tested by two different experiments. The first experiment used pair-wise trials, analyzing the preference by Chi-square tests using two different constructions of the null hypothesis. One hypothesis was based on a no-preference among shell species, the second on comparing the number of crabs changing for a particular shell species when two options were given versus the changing when no options were offered. The second experiment was a multiple-alternative test based on a rank ordering of the shell preference. This method has both statistical and resource-saving advantages over the traditional pair-wise comparisons. The sequence of shell preference was similarly independent of the procedure used. The preferred shell species are heavy and might be associated with hydrodynamic advantages and with the protection against predation. The shell preference matches with the pattern of shell occupancy indicating that the shell use in nature is determined by the crab’s preference. The information generated may be used for further research on shell preference as a methodological alternative.     

Richness,abundance and shell use of subarctic and arctic hermit crabs

Hermit crabs are an obvious, common and abundant feature of global shallow water environments but almost nothing is known of their ecology at their extreme latitudinal ranges, the Arctic and southern South America. In this study, we report the first investigation on hermit crabs and their use of a key resource, gastropod shells, in a subpolar and a polar environment—amongst the most rapidly changing places on earth. Hypothesised low levels of richness were confirmed by surveys—only Pagurus pubescens was found in western Spitsbergen and only three pagurids were found in northern Norway. At the northern-most of their extent, hermit crabs were fairly common but Arctic abundances (1–5 m⁻²) were an order of magnitude lower than in many warm temperate or tropical localities. Along the open coast of Spitsbergen, the occurrence of P. pubescens was infrequent and very patchy, but it was more abundant in the fjords. In Isfjorden, the largest studied fjord, the population of P. pubescens was mainly represented by small individuals. Spitsbergen P. pubescens used few shell types and >87% just occurred in Margarites or Buccinum shells. The proportion of the gastropod shells, used by P. pubescens, which were damaged, was high and increased with shell size. These hermit crabs are at the edge of the range for both their species and Anomura. The extremes of their location are reflected by: They probably represent the least rich assemblage, with the lowest and most patchy typical abundances reported to date and are amongst the smallest individuals using the least diverse and most damaged gastropod shells.     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Influence of size,species and damage on shell selection by the hermit crabPagurus longicarpus

Shell preferences were examined inPagurus longicarpus Say, collected from Wakulla Beach, Florida, USA in 1987. Relative shell size was a more important shell characteristic in shell selection than either shell species or shell damage. Hermit crabs rarely selected relatively large shells but often selected relatively small shells over damaged shells. Preferences for a particular shell species were offset by varying the relative size or condition of the preferred shell species. Because specific environmental pressures are linked to particular shell characteristics, an assessment of the importance of various shell features may indicate which environmental pressures are ameliorated via the gastropod shell. Predation and desiccation are linked to relatively small shells and particular shell species; these stresses may be minimized by alternative behavioral mechanisms.     

Incidence,reproduction and feeding of Stylochus zebra,a polyclad turbellarian symbiont of hermit crabs

The hermit crab Pagurus pollicaris is the common host for the polyclad turbellarian Stylochus zebra in the Atlantic coastal waters from Massachusetts to North Carolina, USA. S. zebra is reported for the first time from two other hermit crabs, Pagurus impressus and Petrochirus diogenes. It was not found with Clibanarius vittatus or Pagurus longicarpus, although the latter serves as a host along the Gulf coast of Florida. The incidence of the polyclad with P. pollicaris in Massachusetts is more than twice that in populations to the south. Multiple infestations (up to 7 worms/crab) occur in over 50% of the infested crabs from Massachusetts, but they are rare elsewhere. Worms living in clear plastic shells with crabs orient in the body whorl dorsal to the host. Egg masses of worms are cemented in the same position within gastropod shells harboring the crabs. In the laboratory, the worm feeds on the gastropod Crepidula plana, a common inhabitant of shells occupied by P. pollicaris. Its feeding behavior is described. The relationship between S. zebra and P. pollicaris appears to be generally commensalistic, but it could be more complicated if the predation of crab embryos observed in the laboratory is confirmed under natural conditions.Contribution No. 755 from the Virginia Institute of Marine Science, Gloucester Point, Virginia, USA.     

Resource degradation: a subtle effect of bottom fishing

Populations of hermit crabs are critically limited by the availability of suitable gastropod shells that they utilise to reduce their risk of predation and environmental stress. Common whelks are the main source of shells for large hermit crabs in the northern Atlantic but are vulnerable to direct and indirect effects of fishing activity. This study examined the potential consequences of degrading shell resources for common hermit crabs. Laboratory trials demonstrated that hermit crabs avoid low-quality damaged shells throughout their life history. This laboratory preference was corroborated by direct field observations of shells preferentially occupied by hermit crabs, compared with shells available for occupation. In the field, 8 times as many empty shells had holes compared to shells occupied by hermit crabs. In the North Sea, the abundance and biomass of live whelks and hermit crabs collected at sites where they co-occurred were significantly related. However, whelks occurred at far fewer sites overall and were more patchily distributed at high abundance than hermit crabs, which were more widespread. At a subset of sites, whelks of the same body-mass range occurred in the Irish and North Sea. However, at these sites, hermit crabs sampled from the North Sea had a significantly lower biomass. This suggests that the shells available for occupation at the North Sea sites would not support crabs of a body mass comparable to that found in the Irish Sea. Using published data, we calculated that in some of the intensively fished areas of the North Sea, 24% of the available shell resource will be damaged each year. The reduction in shell quality in the North Sea may impose a physical constraint on the upper size limit currently attainable by hermit crabs and hence may have implications for population viability.Communicated by J. P. Thorpe, Port Erin     

Distribution of Trypetesa lampas (Cirripedia,Acrothoracica) in various gastropod shells

The distribution of Trypetesa lampas (Hanoock) and its rates of infection of different species of gastropod shells in the waters of Anglesey were studied. T. lampas was found in depths from 3 to at least 25 fathoms (5 to 46 m). It infected dead shells, mainly of Buccinum undatum L., Colus gracilis (da Costa), Neptunea antiqua (L.) and Natica alderi (Forbes); it was absent from shells occupied by hermit crabs in the intertidal zone. Significantly more B. undatum shells were infected than shells of N. antiqua and Natica alderi, and more of both C. gracilis and N. antiqua than of Natica alderi. Larger shells of B. undatum were more likely to be infected than were the smaller ones. The number of females per shell varied with the shell length raised to the power 2.39 instead of as expected with the square of the length.     

Factors affecting the interval between clutches in the hermit crab <Emphasis Type="Italic">Pagurus nigrivittatus</Emphasis>

Interval between reproductive events is an important factor for iteroparous animals because it determines the number of clutches throughout life. This study examined whether female size, clutch size, shell size and prenuptial molting affected the clutch interval in the hermit crab Pagurus nigrivittatus. Precopulatory guarding pairs of P. nigrivittatus were sampled in the field and kept in the laboratory until the female extruded eggs. The clutch interval of each female was assessed as one of two types of relatively “short” and “long” intervals by checking whether the guarded female had eggs and/or egg cases from the preceding brood or not when the guarding pair was collected. The clutch interval was longer in females with prenuptial molting than those without molting and these females usually grew larger at the prenuptial molt. This suggests that female P. nigrivittatus with a long interval might allocate energy into growth at the expense of the number of clutches during the current reproductive season. The allocation to growth is theoretically predicted to decrease with female size. Gastropod shell size is also known to affect the reproductive activity in hermit crabs. However, female size did not significantly affect the clutch interval in P. nigrivittatus, and the effect of gastropod shell size on clutch interval was not consistent with previous empirical studies. These results may be caused by differences in the gastropod species of shell occupied by the females of P. nigrivittatus.