Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

Comparative experimental taphonomy of eight marine arthropods indicates distinct differences in preservation potential

Global biodiversity patterns in deep time can only be understood fully when the relative preservation potential of each clade is known. The relative preservation potential of marine arthropod clades, a diverse and ecologically important component of modern and past ecosystems, is poorly known. We tackled this issue by carrying out a 205‐day long comprehensive, comparative, taphonomic experiment in a laboratory by scoring up to ten taphonomic characters for multiple specimens of seven crustacean and one chelicerate species (two true crabs, one shrimp, one lobster, one hermit crab, one stomatopod, one barnacle and one horseshoe crab). Although the results are preliminary because we used a single experimental setup and algal growth partially hampered observations, some parts of hermit crabs, stomatopods, swimming crabs and barnacles decayed slowly relative to other parts, implying differential preservation potentials within species, largely consistent with the fossil record of these groups. An inferred parasitic isopod, manifested by a bopyriform swelling within a hermit crab carapace, decayed relatively fast. We found limited variation in the decay rate between conspecifics, and we did not observe size‐related trends in decay rate. Conversely, substantial differences in the decay rate between species were seen after c. 50 days, with shrimps and stomatopods decaying fastest, suggesting a relatively low preservation potential, whereas the lobster, calico crabs, horseshoe crabs and barnacles showed relatively slow decay rates, suggesting a higher preservation potential. These results are supported by two modern and fossil record‐based preservation potential metrics that are significantly correlated to decay rate ranks. Furthermore, we speculate that stemward slippage may not be ubiquitous in marine arthropods. Our results imply that diversity studies of true crabs, lobsters, horseshoe crabs and barnacles are more likely to yield patterns that are closer to their true biodiversity patterns than those for stomatopods, shrimps and hermit crabs.     

Land hermit crab (Coenobita clypeatus) densities and patterns of gastropod shell use on small Bahamian islands

Aim To examine patterns of abundance, density, size and shell use in land hermit crabs, Coenobita clypeatus (Herbst), occurring on three groups of small islands, and to determine how these variables change among islands. Location Small islands in the Central Exuma Cays and near Great Exuma, Bahamas. Methods Land hermit crabs were captured in baited pitfall traps and were separately attracted to baits. A mark–recapture technique was used in conjunction with some pitfall traps monitored for three consecutive days. The size of each crab and the type of adopted gastropod shell were recorded, along with physical island variables such as total island area, vegetated area, island perimeter, elevation and distance to the nearest mainland island. Results Relative abundances, densities and sizes of crabs differed significantly among the three island groups. Densities of land hermit crabs were as high as 46 m⁻² of vegetated island area. In simple and multiple linear regressions, the only variable that was a significant predictor of the abundance of hermit crabs was the perimeter to area ratio of the island. Patterns of gastropod shell use varied significantly among the island groups, and the vast majority of adopted shells originated from gastropod species that inhabit the high intertidal and supratidal shorelines of the islands. Main conclusions Although densities of land hermit crabs varied, they were relatively high on many islands, and land hermit crabs may play an important role in these insular food webs. Patterns of shell use may be strongly restricted by island geomorphology: irregular shorelines provide relatively more habitat for the gastropod species that account for the majority of adopted shells and the steep sides of the islands prevent the accumulation of marine gastropod shells. The size of adult hermit crabs appears to be limited by the relatively small gastropod shells available, while the abundance of hermit crabs may be limited by the number of shells available.     

Ecological aspects of hermit crabs (Crustacea,Anomura, Paguroidea) off the northern coast of São Paulo State,Brazil

The occurrence of species of hermit crabs and their ecological distribution in soft bottoms off Ubatuba, São Paulo, Brazil were analyzed. To better understand the distribution of the species in relation to environmental factors, the similarity and species-diversity indexes were calculated. Paguroideans were sampled monthly from January through December 2000. The trawls were made with two otter-trawl nets at 13 different sites, at depths of 2–40 m. Water temperature, salinity, sediment texture, and organic matter content were measured. Gastropod shells occupied by hermit crabs were also assessed. A total of 1,238 specimens was collected, belonging to the families Diogenidae and Paguridae, comprising seven genera and thirteen species. The most abundant hermit crab species were Dardanus insignis (761 specimens) and Loxopagurus loxochelis (351 specimens). Phimochirus holthuisi is newly reported from the São Paulo coast. The highest diversity index was found for the shallower sites near rocky shores. The results of the grouping analysis for sites and species indicated three distinct groups for sites, and four groups for species. This suggests that the occurrence of these anomurans is associated with the environmental and biotic factors analyzed.     

Hermit crab biocoenoses: a worldwide review of the diversity and natural history of hermit crab associates

The symbiotic associates of hermit crabs (excluding parasites and flora) are reviewed worldwide. The review includes species found on the shells occupied by hermit crabs (epibiotic species), species boring into these shells (endolithic species), species living within the lumen of the shell (either free-living or attached to the shell), species attached to the hermit crabs themselves, and hypersymbionts. In total over 550 invertebrates, from 16 phyla are found associated with over 180 species of hermit crabs. Among these associates, 114 appear to be obligate commensals of hermit crabs, 215 are facultative commensals, and 232 are incidental associates. The taxa exhibiting the highest number of associates are arthropods (126), polychaetes (105), and cnidarians (100). The communities of species associated with Dardanus arrosor, Paguristes eremita, Pagurus bernhardus, Pagurus cuanensis, and Pagurus longicarpus are the best studied and harbor the most diverse assemblages of species. While trends in biodiversity of hermit crab assemblages do not follow predicted patterns (e.g., hermit crabs within the Indo-West Pacific do not harbor more species than those from temperate regions), this is suggested to reflect a lack of sampling rather than a true representation of the number of associates. Hermit crabs date to at least the Cretaceous and provided a niche for a number of groups (e.g., hydractinians, bryozoans, polydorids), which were already associates of living gastropods. Apparently hermit crab shells initially supplied a substrate for settlement and then these symbiotic relationships were reinforced by enhanced feeding of symbionts through the activity of the hosts. Through their use and recycling of gastropods shells, hermit crabs are important allogenic ecosystem engineers in marine habitats from the intertidal to the deep sea. Hermit crabs benefit from some symbionts, particularly cnidarians and bryozoans, through extension of shell apertures (alleviating need to switch into new shells) and by providing protection from predators. However, hermit crabs are also negatively impacted (e.g., decreased reproductive success, increased predation) by some symbionts and a review of egg predators is provided. Thus, the symbiotic relationships between hermit crabs and many associates are difficult to characterize and often exhibit temporal changes depending on environmental and biological factors. Research on the biology of these symbionts and the costs/benefits of their associations with hermit crabs are analyzed. While some associates (e.g., Hydractinia spp.) have been studied in considerable detail, for most associations little is known in terms of the impacts of symbionts on hosts, and future experimental studies on the multitude of relationships are suggested.     

Temporal patterns in coral reef, seagrass and mangrove communities from Chengue bay CARICOMP site (Colombia): 1993-2008

Few monitoring programs have simultaneously assessed the dynamics of linked marine ecosystems (coral reefs, seagrass beds and mangroves) to document their temporal and spatial variability. Based on CARICOMP protocol we evaluated permanent stations in coral reefs, seagrass beds and mangroves from 1993 to 2008 in Chengue Bay at the Tayrona Natural Park, Colombian Caribbean. Overall, the studied ecosystems showed a remarkable stability pattern over the monitoring period. While there were annual variations in coral reefs (coral cover) and mangroves (litterfall) caused by hurricane Lenny in 1999, particular trends in seagrass (leaf area index and leaf productivity) appear to reflect the natural variability in this ecosystem. We suggest that monitoring sites at the three marine ecosystems had in general a healthy development in the last 16 years. Our results are critical to locally improve the management strategies (Tayrona Natural Park) and to understand the long-term dynamics of closely associated marine ecosystems in the Caribbean.     

Shell selection behaviour and spatial distribution of three species of intertidal hermit crabs from Hormuz Island,Persian Gulf (Crustacea: Paguroidea)

Shell selection behaviour and spatial distribution of three hermit crab species, Diogenes avarus, D. karwarensis, and Areopaguristes perspicax, were studied at six sites along the intertidal zones of Hormuz Island in the Persian Gulf. 1025 specimens were collected occupying altogether 31 shell species (D. avarus 28 species, A. perspicax 22 species, and D. karwarensis 8 species). Diogenes avarus was found to be by far the most abundant of these three crab species, and Cerithidea cingulata the dominant shell occupied by these hermit crabs. The distribution of the hermit crabs significantly varied (p<0.05) among the sites. The number and the wide diversity of shells occupied in different sites show that the main factor in shell selection for these hermit crabs is the abundance and distribution of shell species in the field.     

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.     

Taphonomy of modern and fossil intertidal gastropod associations from Isla Santa Cruz and Isla Santa Fe, Galápagos Islands

Examination of modern gastropod associations from the low intertidal zone of Isla Santa Cruz suggests that fossil rocky intertidal deposits from this tropical locality will be taphonomically compromised in three ways: (1) Marine hermit crabs, by their use of empty gastropod shells, will mix the shells from varying tidal heights and habitats, thus facilitating mixed associations of such shells in the fossil record, (2) encrusting organisms on crab-inhabited shells are abundant, while boring organisms are almost non-existent, indicating possible differences in postmortem shell retention, and (3) intertidal shells are further taphonomically altered by terrestrial hermit crabs, which transport the shells onto land as well as physically modify the shells. Gastropod fossils from beach and terrace deposits on Isla Santa Fe are interpreted to be a mixed assemblage of rocky intertidal assemblage with few shells indicating influence from marine hermit crabs. Modification of the shell by marine and terrestrial hermit crabs was also evident. A unique polish to the shells at one locality is attributed to the marine iguanas and is only found in the terrace site biologically bulldozed by egg-laying iguanas. Few studies exist on modern rocky intertidal associations in the Galápagos, and the fossil record of rocky shores may provide a baseline for future studies in how community structure has changed over since the advent of humans. Galapagos, C oenobita C ompressus , gastropods, humans, Gulf of California, bionts, nutrients.
Sally E. Walker, Department of Geology, The University of Georgia, Athens, Georgia, USA; 8th September, 1994; revised 28th June, 1995.     

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.     

Scent of death: Evolution from sea to land of an extreme collective attraction to conspecific death

All living organisms must eventually die, though in some cases their death can bring life‐giving opportunities. Few studies, however, have experimentally tested how animals capitalize on conspecific death and why this specialization would evolve. Here, we conducted experiments on the phylogenetically most closely‐related marine and terrestrial hermit crabs to investigate the evolution of responses to death during the sea‐to‐land transition. In the sea, death of both conspecifics and heterospecifics generates unremodeled shells needed by marine hermit crabs. In contrast, on land, terrestrial hermit crabs are specialized to live in architecturally remodeled shells, and the sole opportunity to acquire these essential resources is conspecific death. We experimentally tested these different species’ responsiveness to the scent of conspecific versus heterospecific death, predicting that conspecific death would have special attractive value for the terrestrial species. We found the terrestrial species was overwhelmingly attracted to conspecific death, rapidly approaching and forming social groupings around conspecific death sites that dwarfed those around heterospecific death sites. This differential responsiveness to conspecific versus heterospecific death was absent in marine species. Our results thus reveal that on land a reliance on resources associated exclusively with conspecifics has favored the evolution of an extreme collective attraction to conspecific death.     

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.     

Respiration and Circulation in Land Crabs: Novel Variations on the Marine Design

Both the "true" crabs (Brachyura) and hermit crabs (Anomura)include species that show numerous behavioral, morphological,and physiological specializations permitting terrestrial life.This paper examines respiratory and circulatory adaptationsfor air breathing in these land crabs. Respiratory specializationsinclude modification of gas exchange structures for air breathing(gills and elaborated branchial chamber linings), ventilatorymechanisms permitting effective air pumping, an elevated hemolymphoxygen capacity, and a primarily CO₂- rather than O₂- sensitiveventilatory control system. The qualitative aspects of hemolymphoxygen transport and metabolic rate are apparently unchangedfrom that of marine crabs. While the basic cardiovascular morphologyof land crabs appears similar to that of marine forms, thereis considerable elaboration of the vasculature of the branchialchamber lining, which in some species includes a unique doubleportal system. Cardiac output is lower in land crabs (probablyrelated to their higher hemolymph O₂ capacity), but insufficientdata on hemolymph pressures prevent comparisons with marineforms. In general, land crabs have modified (sometimes extensively)existing structures and processes found in their marine relativesrather than evolving structures for terrestrial life de novo.Accordingly, land crabs present a useful model for the evolutionof terrestriality, showing that even subtle anatomical changescan result in the large changes in physiological function necessaryfor the terrestrial invasion.     

Use of pleopod morphology to determine sexual dimorphism and maturity in hermit crabs: <Emphasis Type="Italic">Isocheles sawayai</Emphasis> as a model

In the Anomura, studies on growth patterns are infrequent, possibly because the heterogeneity of the group, especially in terms of morphology, makes it difficult to construct generalized growth models. Particularly hermit crabs are an interesting group to evaluate aspects of growth, because of their unique body. Isocheles sawayai, a hermit crab found only in the western Atlantic Ocean, poorly known with respect to its sexual dimorphism and maturity, was investigated here based on morphometry. Monthly collections (July 2001 through June 2003) were made from a shrimp fishing boat in the Caraguatatuba region on the northern coast of the state of São Paulo, Brazil. The specimens were measured and weighed, and had their sex checked. Throughout the sampling period, 374 specimens of I. sawayai were collected (11.23% nonovigerous females, 6.69% ovigerous females, 79.41% males and 2.67% intersexes). The size at which morphological sexual maturity was reached by both sexes ranged from 4.0 to 4.3 mm shield length, according to the relative growth and the size of the smallest ovigerous female. Sexual dimorphism was shown by males, which were significantly larger than females, and by differences in growth pattern between the sexes, especially for relationships that involved the pleopods, which is related to their different functions in males and females. The present study is one of the first to use pleopod morphometry to determine sexual maturity and dimorphism in hermit crabs, especially for species with intersexuality such as I. sawayai.     

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to δ¹⁵ N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in δ¹⁵ N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Symbiosis of sea anemones and hermit crabs: different resource utilization patterns in the Aegean Sea

The small-scale distribution and resource utilization patterns of hermit crabs living in symbiosis with sea anemones were investigated in the Aegean Sea. Four hermit crab species, occupying shells of nine gastropod species, were found in symbiosis with the sea anemone Calliactis parasitica. Shell resource utilization patterns varied among hermit crabs, with Dardanus species utilizing a wide variety of shells. The size structure of hermit crab populations also affected shell resource utilization, with small-sized individuals inhabiting a larger variety of shells. Sea anemone utilization patterns varied both among hermit crab species and among residence shells, with larger crabs and shells hosting an increased abundance and biomass of C. parasitica. The examined biometric relationships suggested that small-sized crabs carry, proportionally to their weight, heavier shells and increased anemone biomass than larger ones. Exceptions to the above patterns are related either to local resource availability or to other environmental factors.     

Vacancy Chains Provide Aggregate Benefits To Coenobita clypeatus Hermit Crabs

Vacancy chain theory describes a unique mechanism for the sequential distribution of animal resources across multiple individuals. This theory applies to any resources, such as shelters or nest sites, that are discrete, reusable, and limited in use to single individuals or groups at one time. Hermit crabs rely on gastropod shells for shelter, and a single vacant shell can initiate a chain of sequential shell switches that distributes new resources across many individuals. Using the terrestrial hermit crab Coenobita clypeatus , we examined the previously untested theoretical prediction that this process will yield trickle-down resource benefits to vacancy chain participants (aggregate benefits). In laboratory experiments, we measured improvements in shell quality when a single vacant shell was provided to groups of eight crabs. We found that crabs participating in vacancy chains (averaging 3.2 individuals) gained significant reductions in their shell crowding. In addition, vacancy chains terminated early when experimental groups included a single crab occupying a damaged shell, because damaged vacancies always remained unoccupied. Hermit crabs in damaged shells were more likely to win resource contests for high quality shells against size-matched hermit crabs in crowded shells. Finally, field additions of many new shells to an island population of C. clypeatus hermit crabs reduced average shell crowding for crabs of all sizes, possibly from propagation of benefits through vacancy chains. These results provide empirical support for the theoretical prediction that vacancy chains should provide benefits distributed across many vacancy chain participants. Since shelter-based vacancy chains likely occur in other animals, additional studies of vacancy chain processes should provide new insights into resource acquisition behaviors in diverse animal groups.     

Behavioral Adaptations of Intertidal Hermit Grabs

SYNOPSIS. AS an ecotone, the littoral environment is often complexand is generally rich in numbers and species of organisms. Thedisadvantages of the biotope in terms of exposure to physicalfactors of both the marine and terrestrial environments arepatent, but the advantages are not so evident. The continualreplenishment of food brought from the sea, particularly fordetritus-feeding animals such as hermit crabs, coupled withthe possibility and ability to establish microhabitatswith microclimaticconditions may constitute the principal advantage. Escape fromspecialized predators may also be important. Hermit crabs havesuccessfully exploited most intertidal environments. As membersof the "benthic detritus-feeding guild" food is abundant, andby utilizing their shells in conjunction with movements withinthe littoral zone they have met successfully most of the rigorsof the environment. The shell also provides some protectionfrom predation, particularly from non-specialized predators.Indeed,the behavioral patterns associated with living in shells whichpermit the shell to serve as a microhabitat constitute the majoradaptation enabling the hermit crabs to exploit the intertidalenvironment so successfully.     

The Structure and Density of Aggregations of the Hermit Crab <Emphasis Type="Italic">Pagurus proximus</Emphasis> Komai, 2000 in Epibiosis of the Brown Alga <Emphasis Type="Italic">Costaria costata</Emphasis> (C. Agardh) De A. Saunders, 1895 in the Sublittoral Zone of Vostok Bay,Sea of Japan

A study of epibiosis of the brown alga Costaria costata in the sublittoral zone of Vostok Bay, Sea of Japan during the spring and summer of 2016 showed that hermit crabs of the following four species accounted for approximately 85% of the total biomass: Pagurus proximus, P. minutus, P. brachiomastus, and P. middendorffii. Of these, the most abundant one (96.5% of the total number of hermit crabs) was P. proximus, the mean aggregation density of which on C. costata (429 ± 221 ind./m²) was higher by an order of magnitude than that in the adjacent area of the bouldery bottom. The size ranges of males and females of this species in aggregations on boulders and those on C. costata were similar, with a predominance of larger individuals on algae. In the size–frequency distribution, three groups of individuals corresponding to three size–age cohorts were differentiated. The male-to-female ratio was 1 : 1.7. Hermit crab aggregations on C. costata are ephemeral; their existence is determined by the duration of the period of algae development, that is, from late spring to the middle of the summer.     

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.