Hydrozoan and protozoan epibionts on two decapod species, Liocarcinus depurator (Linnaeus, 1758) and Pilumnus hirtellus (Linnaeus, 1761), from Scotland

Crabs of the species Liocarcinus depurator and Pilumnus hirtellus, collected on the west coast of Scotland, showed hydrozoan and protozoan epibionts. The epibionts of Liocarcinus depurator were the protozoan Ephelota plana and the hydrozoans Clytia gracilis and Leuckartiara sp. The epibionts of Pilumnus hirtellus were the protozoans E. plana and Zoothamnium sp. For Liocarcinus depurator the number of Ephelota per crab fluctuated between 0 and 47 and the greatest number of suctorians were located on the chelipeds, carapace and anterior pereiopods. The hydrozoans, for Liocarcinus depurator, showed densities of 0-20 (Clytia gracilis on the second pereiopod) and 0-507 individuals per crab (Leuckartiara sp., principally on chelipeds, carapace and the fourth right pereiopod). For Pilumnus hirtellus, Ephelota plana showed densities between 3 and 56 individuals per crab, the greatest number of suctorians being located on the same areas as on Liocarcinus depurator. There was a density of 10-69 individuals per crab of Zoothamnium sp. on Pilumnus hirtellus (located on the carapace). Ephelota plana has not been observed previously as an epibiont on crustacea, nor had Zoothamnium sp. as an epibiont on Pilumnus hirtellus. Both hydrozoans, Leukartiara and Clytia, have not been previously described as epibionts on Liocarcinus depurator. Data about the morphological characteristics and distribution of these epibionts are included.     

Invertebrate and protozoan epibionts on the velvet swimming crab Liocarcinus puber (Linnaeus, 1767) from Scotland

Invertebrate and ciliate protozoan epibionts of velvet swimming crabs collected near Millport (Scotland) were analysed. The ecdysis peak for male crabs was at the time of collection while that for female crabs was 2 months later. The epibionts were: the polychaetes Pomatoceros triqueter and Hydroides norvegica, the cirriped Balanus crenatus, the entoproct Barentsia matsushimana, the hydroids Leuckartiara sp. and Clytia sp., and the ciliate protozoans Ephelota plana, Ephelota gemmipara, Chilodochona quennerstedti and Cothurnia longipes. The polychaetes, cirripeds, entoprocts and hydroids, all of them with comparatively larger size, were distributed on the carapace, ventral surface of the cephalothorax and the pereiopods; meanwhile the protozoans, with smaller size, were attached also on pleopods, antennae, eyes and buccal appendages. Chilodochona quennerstedti was the epibiont most abundant on the crab, followed by Ephelota plana. Cothurnia longipes was, in contrast, the least abundant epibiont, followed by Barentsia matsushimana. The anatomical unit most colonized was the left third maxilliped, followed by the left first maxilliped and the ventral surface. The less colonized anatomical units were the left antenna and the ocular orbits. Epibionts on this crab have not been described before. Statistical analyses of the epibiont distribution on the crab were carried out. There were significant correlations between right and left appendages in 66.67% of the cases. Males and females differed significantly with respect to the distribution of epibionts on their anatomical units. The comparison analysis indicated a significant differential distribution of each epibiont species on the anatomical units of the crab. Principal component analysis grouped the epibiont species according to their colonization pattern in three clusters: (1) Ephelota plana, Ephelota gemmipara and Chilodochona quennerstedti; (2) Leuckartiara sp., Clytia sp., Barentsia matsushimana and Cothurnia longipes; and (3) Pomatoceros triqueter, Balanus crenatus and Hydroides norvegica. The hierarchical cluster analysis grouped the anatomical units of the crab in relation to their colonization in five clusters.     

Epibiont protozoan communities on Caridina lanceolata (Crustacea, Decapoda) from the Malili lakes of Sulawesi (Indonesia)

The epibiont protozoan communities living on the freshwater shrimp Caridina lanceolata Woltereck, 1937a from the three major lakes (Towuti, Matano and Mahalona) of the Malili lake system (Sulawesi, Indonesia) were studied. The number of epibionts varied between 2 and 971 per shrimp. Seven protozoan ciliate species were found: Acineta sulawesiensis n. sp., Cothurnia sp., Zoothamnium sp. (in all three lakes), Vorticella sp. (Lake Mahalona and Lake Matano), Opercularia sp. (Lake Mahalona), Epistylis sp. (Lake Mahalona and Lake Matano), and Podophrya sp. (Lake Mahalona). Although these ciliates had been found previously on other crustaceans, they have not been observed as epibionts on Caridina H. Milne Edwards, 1837. The distribution of the different epibiont species on the anatomical units of the shrimp was analyzed in each lake. There is a statistical significant difference between the three lakes in respect to the number of epibionts on each anatomical unit of all analyzed shrimps. The total and mean densities of each epibiont species on the different analyzed shrimps showed a significant difference between the three lakes; i.e., the presence of each epibiont species on the population of C. lanceolata varied from one lake to another. In Lake Towuti the highest density of epibionts was found on the anterior part of the shrimp body (rostrum, antennae, antennulae and eyes) (32.41%), while in the other two lakes, the highest colonization corresponded to the maxillipeds (31.56% Lake Matano, 40.89% Lake Mahalona). In Lake Towuti the rest of epibionts colonized mainly maxillipeds and pleopods (both 45.76% of epibionts). In Lake Matano, other epibionts were distributed principally on the anterior part of the body and pleopods (in total 57.18% of epibionts). In Lake Mahalona, other epibionts were divided among the anterior part of the body, pereiopods and pleopods (in total 57.39% of the epibionts). Uropods and telson were the units less colonized in Lake Matano (3.64%) and Lake Mahalona (1.72%), while in Lake Towuti, they presented a moderate density (13.18% of the epibionts). Taking into account the distribution of epibionts along the antero-posterior axis of the shrimp, considering the different anatomical units, there was a significant correlation between the three lakes. This fact indicates that, in the three lakes, the colonization on C. lanceolata followed a similar distribution pattern, independently of the epibiont species present. The comparison between the distributions of the same epibiont species along the longitudinal axis of the shrimp on the diverse lakes showed that they correlated respect to their density values on the anatomical units of the shrimp. Diverse aspects of the colonization patterns are discussed. Morphological features, taxonomic identification, and particular distribution of the epibiont species in each lake are also included.     

Activities of two British species of Pagurus (Crustacea,Decapoda, Paguroidea)

Two series of experiments were conducted to investigate the activities of two British species of hermit crab, Pagurus bernhardus (L.) and Pagurus prideauxi Leach. The first series showed that P. bernhardus was more active during day than night whereas the opposite was true of P. prideauxi. In the second series, conducted under periods of 24 hours constant illumination, the activity pattern of P. bernhardus persisted to a lesser degree, but P. prideauxi showed no activity pattern. Differences in activity were observed among individuals of the same species, and also between individuals kept in the laboratory for different lengths of time.     

Epibionts of the great spider crab, Hyas araneus (Linnaeus, 1758), in the Barents Sea

Hyas araneus is the most common brachyuran crab in the coastal Barents Sea. Its epibionts were occasionally examined in 1940–1950s. To obtain modern information about associated organisms living on the crabs and compare new data with previous findings, the species composition, infestation indices, and spatial distribution of macro-epibionts colonizing H. araneus were examined in Dalnezelenetskaya Bay, a small semi-open gulf, in summer 2008–2010. A total of 41 taxa were observed on 48 crabs collected from depths ranging from 5 to 28 m. Red algae Ptilota plumosa (prevalence 39.6%) and Palmaria palmata (37.5%) as well as the copepod Harpacticus uniremis (39.6%) and the tube-dwelling worm Placostegus tridentatus (35.4%) predominated on the crabs. Infestation indices were similar in male and female crabs except for the turbellarian worm Peraclistus oophagus. Comparison with the previous study showed some changes in H. araneus fouling community that may be attributed to the shift of climatic regime or differences in sampling procedures. Presence of the amphipod Ischyrocerus commensalis, a new epibiont of H. araneus in the study area may be associated with introduction of the red king crab Paralithodes camtschaticus. Infestation indices of common epibionts depend on the crab shell conditions. Localization of epibionts strongly depended on the settlement patterns of larvae and host-associate relationships. Typical fouling organisms were found predominantly on the carapaces or limbs, while the majority of mobile species were recorded on the gills. Colonization of great spider crabs is beneficial for the epibiont species, whereas some negative effects for the basibiont could not be excluded.     

Comparative brain architecture of the European shore crab <Emphasis Type="Italic">Carcinus maenas</Emphasis> (Brachyura) and the common hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis> (Anomura) with notes on other marine hermit crabs

The European shore crab Carcinus maenas and the common hermit crab Pagurus bernhardus are members of the sister taxa Brachyura and Anomura (together forming the taxon Meiura) respectively. Both species share similar coastal marine habitats and thus are confronted with similar environmental conditions. This study sets out to explore variations of general brain architecture of species that live in seemingly similar habitats but belong to different major malacostracan taxa and to understand possible differences of sensory systems and related brain compartments. We examined the brains of Carcinus maenas, Pagurus bernhardus, and three other hermit crab species with immunohistochemistry against tyrosinated tubulin, f-actin, synaptic proteins, RF-amides and allatostatin. Our comparison showed that their optic neuropils within the eyestalks display strong resemblance in gross morphology as well as in detailed organization, suggesting a rather similar potential of processing visual input. Besides the well-developed visual system, the olfactory neuropils are distinct components in the brain of both C. maenas and P. bernhardus as well as the other hermit crabs, suggesting that close integration of olfactory and visual information may be useful in turbid marine environments with low visibility, as is typical for many habitats such as, e.g., the Baltic and the North Sea. Comparing the shape of the olfactory glomeruli in the anomurans showed some variations, ranging from a wedge shape to an elongate morphology. Furthermore, the tritocerebrum and the organization of the second antennae associated with the tritocerebrum seem to differ markedly in C. maenas and P. bernhardus, indicating better mechanosensory abilities in the latter close to those of other Decapoda with long second antennae, such as Astacida, Homarida, or Achelata. This aspect may also represent an adaptation to the “hermit lifestyle” in which competition for shells is a major aspect of their life history. The shore crab C. maenas, on the other hand seems to rely much less on mechanosensory information mediated by the second antennae but in water, the visual and the olfactory senses seem to be the most important modalities.     

Biodiversity and distribution of epibiontic communities on Caridina ensifera (Crustacea,Decapoda, Atyidae) from Lake Poso: comparison with another ancient lake system of Sulawesi (Indonesia)

Fernandez‐Leborans, G. and von Rintelen, K. 2010. Biodiversity and distribution of epibiontic communities on Caridina ensifera (Crustacea, Decapoda, Atyidae) from Lake Poso: comparison with another ancient lake system of Sulawesi (Indonesia). — Acta Zoologica (Stockholm) 91 : 163–175 The epibiont communities of the shrimp Caridina ensifera, endemic to Lake Poso (Sulawesi, Indonesia), were analysed. Most of the epibiont species were ciliated protozoa belonging to three suctorian genera (Acineta, Podophrya and Spelaeophrya), three peritrich genera (Zoothamnium, Vorticella and Cothurnia), and a haptorid genus (Amphileptus). There was also a rotifer epibiont of the genus Embata. Epibionts were identified to species level. There were 14 to 1114 epibionts per shrimp. The distribution of the epibiont species on the surface of the basibiont was recorded, calculating the number on the different colonized individuals of C. ensifera. The most abundant species, Zoothamnium intermedium and Acineta sulawesiensis, were also the most widely distributed. There was a significant difference between the spatial distributions of the different epibiont species. The analysis of the number of the epibiont species throughout the anteroposterior axis of the shrimp showed a gradient from the anterior to the posterior end of the body. Data from Lake Poso were compared with those of the Malili lake system (Sulawesi), obtained from its endemic shrimp, Caridina lanceolata. Lake Poso had the highest mean diversity, while Lake Mahalona showed the highest maximum diversity. All lakes were correlated with respect to the mean number of epibionts on the anatomical units of the shrimp, which showed a similar general distribution. The distributions of the different epibiont species were compared between the lakes. The possible adaptations of the epibionts as well as the colonization patterns were discussed. From the statistical results and the analysis of the distributions, we propose that in these communities epibiont species have a pattern of colonization in which they follow a behaviour as a whole; each species has a differential distribution, with the species occupying the available substratum with the particular requirements of each functional group, but there is a trend towards maintaining an equilibrium among species and groups, compensating for diversity and number of individuals. In all lakes there was an epibiont distribution model comprising the maintenance of an anteroposterior axis gradient, which was supported by the fluctuation in diversity and number of individuals of the different functional groups of epibiont species. The functional role of the different groups of species seems to tend towards sustainability with little global variation among the lakes.     

Filter feeding in the hermit crab

Summary The hermit crab, Pagurus bernhardus is able to remove both Artemia nauplii and unicellular algae from suspension. Crabs with wet weights of 1.1–9.2 g consumed all of the 300 Artemia nauplii contained in 200 ml of sea water within 1 h. Crabs weighing 0.7–1.1 g wet weight filtered suspended Dunaliella algae (8 m) from concentrations of 10–350 million cells per liter at a rate of 26% and 47% within 2 and 5 h, respectively. A similar result was obtained with an 11 g crab. During filter feeding activity a water current is generated by the flagella of the exopods of the second and third maxillipeds. Artemia nauplii are caught by grasping movements of the endopods of the third maxillipeds, whereas filtering of unicellular algae is probably achieved by the two maxillae. It is assumed that filter feeding activity depends on the same structures and behavior as in deposit feeding. P. bernhardus is one more example of a benthic marine animal which may use any food source which becomes available in the course of the seasons.     

Associational resistance of fouled blue mussels (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) against starfish (<Emphasis Type="Italic">Asterias rubens</Emphasis>) predation: relative importance of structural and chemical properties of the epibionts

Several epibiotic species reduce starfish (Asterias rubens) preference for the blue mussel Mytilus edulis in the Baltic. The aim of this study was to reveal whether this associational resistance was caused by structural or chemical aspects of the different epibionts. To assess structural epibiont effects, an in situ experiment was conducted with unfouled mussels and mussels equipped with artificial epibionts (dummies) exposed to natural predation by A. rubens. The chemically inert dummies closely matched the structural properties of the locally common epibionts Balanus improvisus (barnacle), Ceramium strictum (red alga), Halichondria panicea (sponge), and Laomedea flexuosa (hydrozoan). Starfish fed indiscriminately in all treatments. Chemical effects of epibionts on the attractiveness of mussels for A. rubens were investigated by incorporating freeze-dried epibionts or mussel tissue into Phytagel pellets at natural concentrations. Starfish were allowed to choose among these structurally similar but chemically different prey items in an in vitro experiment. The predators exhibited significant preferences among the food pellets, which closely matched their preferences for corresponding natural mussel–epibiont associations. Thus, chemical aspects of epibionts appear to play a larger role in this associational resistance than do structural aspects. Implications of these indirect interactions for benthic communities are discussed.Communicated by H.-D. Franke     

Epibiotic protozoan communities on juvenile southern king crabs (<Emphasis Type="Italic">Lithodes santolla</Emphasis>) from subantarctic areas

In this study, we describe for the first time the composition of epibiotic protozoan communities on juvenile southern king crabs Lithodes santolla. Basibionts were collected in subantarctic bays near Santa Ana Point (Magellan Strait, Chile). Seven epibiotic protozoan ciliates were found: Ephelota gemmipara, Ephelota gigantea, Podophrya fixa, Acineta tuberosa, Zoothamnium duplicatum, Chilodochona quennerstedti and Gymnodinioides sp. The mean number of epibionts per crab was 99 (maximum 897). Both Ephelota species were the most abundant and most widely distributed epibionts on the crab examined. The lengths of basibionts were correlated with the number of epibionts, indicating a relationship between the age of the crab and the state of epibiosis. Epibionts differed in their distribution patterns along the anteroposterior axis of the crab. Less abundant species tended to occupy sites that are not colonized by the predominant Ephelota species. Maxillipeds and pereiopods were most densely colonized by epibionts. The behaviours of epibionts and basibionts related to epibiosis are discussed. The recent studies about invasive lithodids in Antarctic areas enhance the interest for the epibiotic communities colonizing these crustaceans, which can illustrate changes in biodiversity and state of these environments.     

An analysis of inter‐ and intraspecific aggregations in two sympatric species of hermit crab (Decapoda,Anomura, Paguridae)

A series of experiments was conducted on the relationship between population density and spacing out in two sympatric species of crab, Pagurus bernhardus (L.) and Pagurus prideauxi Leach. A second series of experiments investigated the two species’ aggressiveness and activity by observing the behaviour of pairs of animals.

In the first series of experiments, animals became more aggregated (Clarke and Evans’ R) as the population density (p) was increased. The rate of aggregation (dR/dp) was greater when the species were mixed than when they were separate.

In the second series of experiments, activity, total encounters, and resolved encounters, were highest in paired P. bernhardus, lower in mixed pairs of the two species, and lowest in pairs of P. prideauxi. Larger individuals were more active than smaller ones in P. bernhardus and P. prideauxi pairs. In the mixed pairs, P. bernhardus were more active. Paired P. bernhardus were slightly more active than isolated P. bernhardus but this difference was not statistically significant. The activities of paired and isolated P. prideauxi were not significantly different. The distances between individuals in pairs were similar for all combinations.

The results are discussed in relation to the distribution of sympatric species.     

Shelter association between the hermit crab Sympagurus dimorphus and the zoanthid Epizoanthus paguricola in the southwestern Atlantic Ocean

Schejter, L. and Mantelatto, F.L. 2011. Shelter association between the hermit crab Sympagurus dimorphus and the zoanthid Epizoanthus paguricola in the southwestern Atlantic Ocean. —Acta Zoologica (Stockholm) 92 : 141–149. The available literature on zoanthid–hermit crab associations deals only with records of this phenomenon, providing no detailed information. We describe, for the first time, the shell‐like colonies of Epizoanthus paguricola associated with the hermit crab Sympagurus dimorphus from benthic samples taken in the Argentine Sea, between 85 and 131 m depth, and provide information about morphometric relationships between the hermits and the zoanthids. In total, 260 specimens (137 males and 123 females) of S. dimorphus were collected, 240 (92.3%) of which were living in symbiosis with E. paguricola. The remaining 20 (7.7%) were living inside gastropod shells. As the initial structure of the pseudoshell, 12 different gastropod species were found (all were almost totally covered with colonies of E. paguricola). The hermit crab lives in the spiral cavity inside the soft colony, which seemed to be slightly different depending on the initial gastropod. Aperture pseudoshell morphology did not seem to be related to the sex of the hermit crab host, although males showed larger apertures for a given colony size. This fact is probably related to a larger size of male’s cheliped (sexual dimorphic character) used like a gastropod operculum and that may serve as a template for the growing of the aperture pseudoshell edge. The number of epizoanthid polyps per colony increased in relation to the weight of the colony and to the size of the hermit crab. A process of selection of the initial shell was evident, because species of Naticidae were not the most common gastropods in this benthic community, but were those most used by hermit crabs (>60%). The puzzling association between hermit crab, shell and zoanthid presumably occurs during the hermit juvenile phase, when the crab occupies a small shell, and a zoanthid larva settles on it. Given the close relationship between S. dimorphus and E. paguricola found in this region, we support the idea that due to the low availability of adequate gastropod shells for hermit life cycle, this association allows the establishment and the continuity of the hermit crab population in the studied area.     

Hermit crabs, humans and Mozambique mangroves

There is a complex interrelationship between upper shore hermit crabs (such as Coenobita sp. and Clibanarius sp.), coastal human populations and mangrove forests in Mozambique. The abundance, activity, shell selection and behaviour of three species of hermit crab are related to the level of mangrove cover. With increased density of mangrove trees, the study species of hermit crab changed in abundance, tended to become diurnal, spent more time feeding and were clustered in larger groups when doing so, and selected longer spired shells. All five of the same variables are also linked to the proximity and activity of humans through both direct and indirect actions. Direct effects included a tendency to nocturnal activity with proximity to human activity; indirect effects included increased and more clumped food supplies, and shell middens from intertidal harvesting and deforestation. Mangroves are important to local human populations as well as to hermit crabs, for a wide variety of (similar) reasons. Mangroves provide storm shelter, fisheries and fishery nursery grounds for adjacent human settlements, but they also harbour mosquito populations and their removal provides valuable building materials and fuel. Hermit crabs may be useful (indirectly) to coastal human populations by being a source of food to certain commercial species, and by quickly consuming rotting/discarded food and faeces (thereby reducing disease and pests). They can also cause minor problems to coastal human populations because they use shells of (fisheries) target mollusc species and can be more abundant than the living molluscs, thereby slowing down effective hand collection through confusion over identification. The mixture of positive and negative attributes that the three groups impart to each other in the Quirimba Archipelago, northern Mozambique, is discussed.     

Molecular evidence of digestion and absorption of epibiotic bacterial community by deep-sea crab Shinkaia crosnieri

The hydrothermal vent crab Shinkaia crosnieri is considered to obtain nutrition from the epibiotic bacteria found on the setae, but previous studies have not shown how nutrients can be transferred from the epibionts to the host. In this study, microscopic observations of S. crosnieri intestinal components detected autofluorescent setae fragments and pigmentation derived from the digestion of epibionts in a dye-stained epibiont tracer experiment. An in vitro digestion experiment with epibiotic populations using an intestinal extract demonstrated the degradation of epibiotic cells by digestive enzymes. A phylogenetic analysis showed that many of the bacterial 16S ribosomal RNA gene sequences obtained from the intestine were closely related to the sequences of the epibionts, thus they were probably derived from the epibionts. A stable isotope tracer experiment also indicated that ¹³C assimilated by the epibionts provided a carbon (nutrition) source for the host. Both activity measurements and isotope studies showed that chemosynthetic metabolism by the gut microbial components were inactive. Together with the feeding behaviour of living S. crosnieri, these results indicate that S. crosnieri ingests the epibionts using maxillipeds and assimilates them via its digestive organs as a nutrient source. The results of this study elucidate the mechanism of nutritional transfer in ectosymbiosis between chemosynthetic bacteria and deep-sea invertebrates.     

Shell occupancy of the intertidal hermit crab Clibanarius erythropus (Decapoda,Diogenidae) on São Miguel (Azores)

The intertidal hermit crab Clibanarius erythropus was collected at three sites on São Miguel (Azores) during low spring tides. Shells occupied were identified and measured. Crab sizes ranged from 1.78 to 13.67 mm (cephalothoracic shield length), with an average size of 4.40 ± 1.44 mm. Of the 19 different shells utilised, the most frequent were Littorina striata (23.8%), Nassarius incrassatus (22.5%) and Mitra sp. (22.0%). At Fenais da Luz, L. striata was most frequently occupied, while at Água de Alto it was N. incrassatus and, at Caloura, Mitra sp. shells were most frequently used. Shell selection appears to be determined by respective sizes of hermit crab and shell species. Small size-class crabs occupy more shell species than larger crabs. The smallest crab was found at Fenais da Luz occupying a small Bittium sp., whereas the largest crab was found at Caloura inhabiting Stramonita haemastoma.     

COLONIZATION AND REPRODUCTION OF THE EPIBIOTIC FLAGELLATE COLACIUM VESICULOSUM (EUGLENOPHYCEAE) ON DAPHNIA PULEX1

The epibiotic flagellate Colacium vesiculosum Pringsheim attaches to planktonic species of Daphnia in freshwater habitats. Previous studies found that prevalence (percentage of substrate organisms carrying attached epibionts) and intensity (number of attached epibionts on a given substrate organism) are low early in the Daphnia intermolt period and are high late in the intermolt period. We tested the hypothesis that increases of Colacium cells attached to Daphnia occur both by rapid initial and continuous colonization and by cell reproduction. Epibiont prevalence and intensities were determined at successive intermolt stages of Daphnia pulex Leydig collected from freshwater ponds in Colorado. Colonization was continuous throughout the intermolt period and was most important to epibiont population increase at the beginning of the intermolt period. Cell division was the major contributor to epibiont increase at the end of the intermolt period.     

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.     

The ecology of Belizean mangrove-root fouling communities: patterns of epibiont distribution and abundance,and effects on root growth

The aerial prop roots of the neotropical red mangrove,Rhizophora mangle L., begin growing well above highest high water (HHW) and often extend well below lowest low water (LLW) before rooting in the benthic substratum. In Belize, Central America, prop roots growing below LLW are colonized by diverse assemblages of organisms, including macroalgae, hydrozoans, ascidians, sponges, anemones, hard corals, and isopod crustaceans. Mangroves, root-fouling epibionts, root herbivores, and benthic predators engage in complex interactions that are major determinants of mangrove growth and production. Species richness of root epibionts increases with distance from the mainland and with proximity to the barrier reef. Species richness decreases with variability in water temperature and salinity. Ascidians and sponges transplanted from Lark Cay into the coastal Placencia Lagoon failed to survive, but anemones from Lark Cay survived in Placencia Lagoon. Reciprocal transplants survived off-shore. The gastropod predator,Melongena melongena L., present only in mainland estuaries, reduced local barnacle abundance and epibiont species richness in Placencia Lagoon. Isopod species richness also increases with distance from shore, but the number of roots bored by these species decreases. These isopods can reduce root relative growth rate (RGR_root) by 55%. On off-shore cays, sponges and ascidians ameliorate negative effects of isopods. In mainland estuaries where epibionts are less common, isopod damage to roots is more severe. Experimental studies in mangrove swamps throughout the world would clarify the importance of plant-animal interactions in these widespread tropical ecosystems.     

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.