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     

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.     

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     

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     

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.     

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     

Metabolic consequences of shell choice in Pagurus bernhardus: do hermit crabs prefer cryptic or portable shells?

Hermit crabs are unique among crustaceans in that they choose their shell and can, therefore, be used to investigate apparent preferences for the presence of epizoic fauna. These may be beneficial due to increased protection from predators via enhanced crypsis but may also make the shell more difficult to carry. The effects of encrusting fauna on the physical properties of empty gastropod shells, the shell preferences of hermit crabs and the physiological costs of shell carrying were examined in the present laboratory study. Heavily encrusted shells weighed more than shells of the same internal volume that were free of epizoic fauna, hermit crabs showed a clear preference for un-encrusted shells and those that occupied encrusted shells had elevated haemolymph lactate in comparison to those supplied with un-encrusted shells. Under laboratory conditions, any benefit from increased crypsis is, therefore, outbalanced by the increased costs of carrying encrusted shells.     

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.     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

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.     

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.     

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     

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.     

Ecology of subtropical hermit crabs in S.W. Madagascar: cluster structure and function

The Madagascar coast (both) has a higher density and diversity of hermit crabs than is known from any other locality in the western Indian Ocean. Of the 20 species occurring at Anakao (S.W. Madagascar), 11 aggregated into clusters, including all but one of the species above the subtidal zone. The mean number of hermit crabs and species in clusters varied with several spatial parameters and time. Over 80% of the community clustered in certain habitats in particular tidal zones, whilst as low as 3% clustered in others. The highest intensity of clustering with shore zone was coincident with peak numbers of hermit crabs. The initiation and duration of clusters of hermit crabs above the eulittoral was driven by circadian rhythms, whilst those in the eulittoral were governed by tidal state. Clusters above and below the eulittoral were longer in duration, and those in the subtidal were more temporally variable than those above it. Certain pairs of species showed positive correlations of occurrence and (more rarely) of abundance, and all the correlations (of occurrence) of one, Clibanarius eurysternus, were negative. Positive correlations of occurrence were related to the degree of shell-use commonality between species pairs. Eulittoral species clustered with other individuals of approximately similar size and exchanged shells upon cluster disintegration. There was evidence of a dehydration-reduction function to shell clustering in addition to shell-exchange facilitation. The highly variable species-specific strategies of clustering may be important in alleviating both intra- and interspecific competition in assemblages of similar and highly abundant species.     

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     

Genetic differentiation between morphotypes of the marine sponge Suberites ficus (Demospongiae: Hadromerida)

Sponges of three morphotypes of Suberites ficus (Johnston, 1842) were collected during February and March 1985 off the south-west of the Isle of Man, and were compared by using spicule size distributions and genetic allele frequencies of isozyme loci. The populations did not show any significant differences of spicule size or type, but could be easily differentiated into three separate species based on isozyme patterns. Samples of pale orange S. ficus growing on gastropod shells inhabited by hermit crabs (Pagurus spp.) were reproductively isolated from the redorange and the pale yellow colour morphs encrusting the bivalve Chlamys opercularis. These latter two colour morphs were genetically similar, but significant differences were observed at two of the 19 gene loci assayed. All the sponges studied were sympatric, and therefore the genetic differences, indicating reproductive isolation, are strong evidence for separate gene pools and, hence, that they are different species. The genetic identity between the two colour morphs of S. ficus on C. opercularis shells was 0.977, whilst between each of these and S. ficus on hermit crabs it was about 0.65. In all three species genetic variability was high, with mean expected and observed heterozygosity values per locus ranging from 0.17 to 0.36.     

Comparing community structure on shells of the abalone Haliotis midae and adjacent rock: implications for biodiversity

This paper concerns the effects on biodiversity of depletion of the South African abalone Haliotis midae, which is a long-lived species with a large corrugated shell that provides a habitat for diverse benthic organisms. We compared community structure on H. midae shells with that on adjacent rock at three sites (Cape Point and Danger Point sites A and B) and at two different times of the year at one of these sites. Shells of H. midae consistently supported communities that were distinctly different from those on rock. In particular, three species of non-geniculate (encrusting) corallines, Titanoderma polycephalum, Mesophyllum engelhartii and Spongites discoideus, were all found either exclusively or predominantly on shells, whereas another non-geniculate coralline, Heydrichia woelkerlingii, occurred almost exclusively on adjacent rock. The primary rocky substratum, however, supported a higher number of species than abalone shells. Possible reasons for the differences between the two substrata include the relative age, microtopography and hardness of the substrata; the abundance of grazers on them; and the relative age of different zones of the abalone shell, which support communities at different stages of succession. Diversity on shells was lowest in zones that were either very young or very old, in keeping with the intermediate disturbance hypothesis. The distinctiveness of shell epibiota will increase β diversity despite having a lower α diversity than that of adjacent rock. Decimation of H. midae by overfishing therefore has implications for biodiversity conservation.     

Epibiotic community of the horseshoe crab Tachypleus gigas

Horseshoe crabs act as moving substrata for simple to complex communities of small marine organisms. Amplexed adult pairs migrate for breeding once every 2 weeks from deep waters towards nearshore waters during highest high tide. Female horseshoe crabs bury themselves to the level of the lateral eyes to deposit eggs while the male crabs fertilize them. Subsequently eggs are buried by the female. Tachypleus gigas (Müller) is the most abundant horseshoe crab species above available along the Orissa coast (India). Adults reach terminal anecdysis once sexually mature and live with their carapace for 4 to 9 years. In spite of this, epibiosis is limited. In the current investigation, differences in the epibiotic community (diatoms and macro-epibionts) present on horseshoe crabs, according to gender, were evaluated, and the macro-epibiont population from different regions of the carapace was mapped. In general, female horseshoe crabs harbored fewer epibionts than the males. Among the diatoms, Navicula spp., Nitzschia spp. and Skeletonema sp. were dominant in both sexes. However, the abundance and diversity of diatoms was greater on the carapaces of male crabs. Among the macro-epibionts, the acorn barnacle (Balanus amphitrite Darwin) and encrusting bryozoan (Membranipora sp.) were the most dominant forms. Barnacles and bryozoans were greater in abundance in the “rough” zone (cardiopthalmic region and anterior region of the opisthosoma). Mapping of the macro-epibionts from different regions of the carapace revealed differential distribution in males and females. Such differentiated distribution of the macro-epibionts can be related to factors such as changing habitat by the horseshoe crabs during breeding, mechanical abrasion and surface availability during mating and nesting periods, requirements of epizootic larvae and surface properties of the carapace (wettability and roughness). In the case of females, mechanical abrasion and surface availability played an important role in the epibiotic community structure and distribution patterns. The surface wettability measurements indicated male carapace to be slightly more hydrophobic than the female carapace. Scanning electron microscopy revealed that the male carapace was comparatively rough compared to the smooth carapace of females. A comparison of surface properties of the carapace indicated that the male carapace is more conducive for epibiosis. Received: 23 August 1999 / Accepted: 25 January 2000     

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.     

Mating success of male hermit crabs in shell generalist and shell specialist species

Summary The reproductive behavior of two species of diogenid hermit crabs was studied in Hawaii. In the shell generalist, Clibanarius zebra, male reproductive success varied little with size, although the largest males were less successful in obtaining copulations than were medium-large males. Male and female size were positively correlated, in successful pairs, thus larger males had the potential to fertilize more eggs when they were successful in obtaining a copulation. Female fecundity in C. zebra was not affected by species of gastropod shell inhabited once female size was taken into account. Male copulatory success was very strongly influenced by the species of gastropod shell inhabited. Males in Trochus or Nerita shells had greatly reduced reproductive success compared to males in Turbo or Nassarius shells. This result was due both to (1) males in Trochus especially dropping and otherwise poorly handling females during precopulatory behavior and (2) females not responding to precopulatory behavior patterns executed by males in Trochus and Nerita. Transferring males from good to bad shapes of shells and vice versa showed that male success was a function of shell type inhabited and not some correlated feature of the crabs. In the shell specialist, Calcinus seurati, which is found primarily in Nerita shells as an adult, males in Nerita shells were quite successful in obtaining copulations.