Hitching a ride on a hermit crabs home: Movement of gastropod shells inhabited by hermit crabs

There is little information on the movement and connectivity patterns for many species. The movement by shells occupied by the hermit crab Pagurus bernhardus and the organisms encrusting these shells was investigated on the south coast of Wales (UK). Hermit crabs shells moved considerable distances along the shore over 1 month periods, moving a minimum mean distance (±SE) of 148 (±8) and 174 (±9) m from release sites in the January and August respectively. Hermit crab-inhabited shells were also found to travel across habitats (sandy areas) that are unsuitable for both hermit crabs and epibionts. Through the examination of 150 of the most abundant shells used by P. bernhardus (Nucella lapillus), twelve epibiont species were found (10 animal and 2 algal species) and choice experiments demonstrated that hermit crabs preferred epibiont covered shells (84%), compared with bare shells (16%). The distance that shells were moved and the preference of hermit crabs for epibiont encrusted shells, coupled with the ability for epibionts to cross unsuitable habitats, may provide a dispersal advantage for epibiont species.     

Indirect Effects of Epibiosis on Host Mortality: Seastar Predation on Differently Fouled Mussels

In situ experiments were run with the seastar Asterias rubens to investigate the influence of epibiosis on predation preferences. Mussels ( Mytilus edulis ) monospecifically fouled by different epibiont species (the barnacle Balanus improvisus , the red filamentous alga Ceramium strictum , the sponge Halichondria panicea and the hydrozoan Laomedea flexuosa ) and macroscopically clean mussels were exposed and seastar predation was monitored by SCUBA. Asterias rubens preferred macroscopical unfouled mussels as prey. Fouling generally reduced predation pressure on the mussel hosts (associational resistance). Barnacles protected mussels less efficiently than hydrozoans or algae.
We hypothesize that in top-down controlled communities this influence of epibiosis on predation pressure should affect mussel community patterns. A survey of natural mussel-epibiont distribution in the presence or absence of A. rubens showed that the prevalence of differently fouled mussels differed between predation-exposed and predation-protected habitats. Natural mussel-epibiont associations reflected the preferential predation of the major local predators. Additionally, higher epibiotic diversity and evenness could be observed at locations accessible to benthic predators as compared with habitats protected from predation.
As blue mussels and seastars are important structuring and controlling elements in the shallow water community of Kiel Fjord, major consequences of epibiosis on the entire system are discussed.     

Differential host use affects fecundity of the gastropod Crepidula onyx

For epibiotic or symbiotic marine invertebrates, alternative host species may differ substantially in quality, and under some circumstances such differences in host quality may lead to the evolution of increased host specificity. However, the fitness consequences of alternative hosts for epibiotic or symbiotic marine invertebrates have rarely been quantified. In Southern California, the gastropod Crepidula onyx is often found as an epibiont on either bay mussels (Mytilus galloprovincialis) or cone snails (Conus californicus). These hosts differ greatly in maximum size, with possible effects on size at sex change and final size in Cr. onyx, and thus on fecundity. Further, Cr. onyx on the two hosts differ in shell shape, possibly affecting the size of the ctenidium, which Cr. onyx uses for suspension feeding. We examined these potential effects of host use on fitness components in Cr. onyx. Epibionts on mussels reached much larger average sizes than did those on cone snails; further, epibionts on mussels often completed sex change at much larger sizes than did those on cone snails. On average, mussel epibionts had threefold higher average fecundities than did cone snail epibionts. Although there was a slight difference in shell shape between epibionts on the two host species, there was no difference in the scaling of ctenidium area with body size for Cr. onyx from the two hosts. The large average differences in fecundity in epibionts associated with the two alternative hosts suggests that there may be strong selection on host choice at larval settlement.     

16S rRNA is a better choice than COI for DNA barcoding hydrozoans in the coastal waters of China

Identification of hydrozoan species is challenging, even for taxonomic experts, due to the scarcity of distinct morphological characters and phenotypic plasticity. DNA barcoding provides an efficient method for species identification, however, the choice between mitochondrial cytochrome c oxidase subunit I（COI） and large subunit ribosomal RNA gene（16S） as a standard barcode for hydrozoans is subject to debate. Herein, we directly compared the barcode potential of COI and 16S in hydrozoans using 339 sequences from 47 pelagic hydrozoan species. Analysis of Kimura 2-parameter genetic distances（K2P） documented the mean intraspecific/interspecific variation for COI and 16S to be 0.004/0.204 and 0.003/0.223, respectively. An obvious ＂barcoding gap＂ was detected for all species in both markers and all individuals of a species clustered together in both the COI and 16S trees. These results suggested that the species within the studied taxa can be efficiently and accurately identified by COI and 16S. Furthermore, our results confirmed that 16S was a better phylogenetic marker for hydrozoans at the genus level, and in some cases at the family level. Considering the resolution and effectiveness for barcoding and phylogenetic analyses of Hydrozoa, we strongly recommend 16S as the standard barcode for hydrozoans.     

Interpreting odours in hermit crabs: A comparative study

Odours of different sources can indicate to hermit crabs the availability of empty shells, crucial resources for the life cycle of almost all of them. Here, we compared Clibanarius erythropus and Pagurus bernhardus for the intensity of investigative behaviour exhibited towards an empty, well-fitting shell in the presence of (1) plain seawater as control and seawater conditioned by (2) dead and live snails, (3) dead and live conspecifics, (4) live predators, and (5) food. During 10 min of observation, we recorded latency (the time until the first contact with the shell), and the number and duration of shell investigation bouts. The two species behaved similarly when exposed to the odours of food, live snails, and predators, while a more intense shell investigation was induced by dead snail odour in C. erythropus and by dead or live conspecific odour in P. bernhardus. Further studies should investigate the influence of phylogeny and ecology on this interspecific difference.     

Associational resistance mediates predator--prey interactions in a marine subtidal system

There is a growing awareness of the role that indirect interactions play in influencing food webs and ecosystem structure. In this study, the hypothesis that crustose algal epibionts provide gastropods associational resistance from predation was investigated through field surveys and laboratory feeding assays. In rocky low intertidal/shallow subtidal systems in the northeast Pacific, several species of crustose algae (the red alga Peyssonnelia meridionalis and crustose corallines) can colonize the shells of living Tegula brunnea snails. The growth patterns of these epibiontic crustose algae allow them to cover their host’s surface completely, which may, in turn, protect their hosts from predation. A multi‐site field survey of T. brunnea revealed that >60% of snails were at least 75% covered with one or more species of crustose algae, with 35% fully covered, indicating that this is common in the field. Laboratory feeding assays revealed that sea stars, a primary predator of T. brunnea, distinguished among snails with different shell coverings; Pisaster consumed nearly three times as many bare (i.e. no crustose algae) snails as those covered with Peyssonnelia, while Pycnopodia consumed four times as many bare snails as those covered with crustose corallines. These results suggest that epibiont crustose algae can benefit their hosts via associational resistance; this finding may have implications for the role of associational resistance in trophic interactions.     

The role of two sediment-dwelling invertebrates on the mercury transfer from sediments to the estuarine trophic web

The annual total and organic mercury bioaccumulation pattern of Scrobicularia plana and Hediste diversicolor was assessed to evaluate the potential mercury transfer from contaminated sediments to estuarine food webs. S. plana was found to accumulate more total and organic mercury than H. diversicolor, up to 0.79 mg kg⁻¹ and 0.15 mg kg⁻¹ (wet weight) respectively, with a maximum annual uptake of 0.21 mg kg⁻¹ y⁻¹, while for methylmercury the annual accumulation was similar between species and never exceeded 0.045 mg kg⁻¹ y⁻¹. The higher organic mercury fraction in H. diversicolor is related to the omnivorous diet of this species. Both species increase methylmercury exposure by burrowing activities and uptake in anoxic, methylmercury rich sediment layers. Integration with the annual biological production of each species revealed mercury incorporation rates that reached 28 μg m⁻² y⁻¹, and to extract as much as 11.5 g Hg y⁻¹ (of which 95% associated with S. plana) in the 0.4 km² of the most contaminated area, that can be transferred to higher trophic levels. S. plana is therefore an essential vector in the mercury biomagnification processes, through uptake from contaminated sediments and, by predation, to transfer it to economically important and exploited estuarine species.     

Shell damage and shell repair in the Antarctic limpet Nacella concinna from King George Island

Nacella concinna is the most conspicuous macroinvertebrate in the intertidal of King George Island. An important predator, the Kelp gull Larus dominicanus, feeds on Nacella during spring low tides. The gulls deposit empty Nacella shells as regurgitates mainly on roosts on coastal rocks. The regurgitates were found to consist of 40% shell fragments by weight and 60% intact shells. Faeces of Kelp gulls contained much smaller fragments than the regurgitates. Some of the Nacella, particularly those too large to ingest, are handled in the intertidal. The middens are, therefore, inadequate to study size selection by Kelp gulls: the largest Nacella are underrepresented. Seventy-five per cent of the intact Nacella shells from the Larus middens showed one or more shell repairs. Such repairs may be due to unsuccessful attacks by gulls, but more probably they indicate damage caused by rolling ice blocks and stones in the intertidal and shallow subtidal. A number of living Nacella were found stranded on the beach, detached from the rocks. They showed damage along the shell margin and even one Nacella was collected without any shell left. The observed repair frequency of 75% in Nacella was much higher than in other (smaller) intertidal gastropods at Potter Peninsula (3–11%, av. 8%). Comparably high frequencies are observed for instance in tropical intertidal gastropods, where repair is due to heavy unsuccessful crab predation; however, shell-crushing crabs are absent on King George Island. This indicates that palaeontologists should be cautious in ascribing all shell repairs in fossil shells (particularly from tidal environments) to predators. Shell repair in the related Nacella deaurata, collected in a less exposed site at Port Stanley (Falkland Islands), occurred only in 13% of the specimens. Another conspicuous form of shell damage was due to grazing by Nacella on the boring algae living in other Nacella shells. Epigrowth of crustose calcareous algae inhibited such grazing, but in the absence of epigrowth deep hollows were scraped in the shells, the parallel scratches by the radula clearly visible, urging Nacella to repair its shell by producing more shelly material on the inside.     

Algal epibiosis on Megabalanus tintinnabulum and its role in segregation of the Balanus amphitrite population

Ulva sp. and Gracilaria sp. were found colonizing shell surfaces of an acorn barnacle, Megabalanus tintinnabulum. However, this association was not noticed in the case of Balanus amphitrite, which was a co‐inhabitant. Such a difference in intergeneric interactions with the algae was examined in laboratory experiments. For this, the influence that extracts of algae, extracts of algae‐associated bacteria and natural leachants from M. tintinnabulum exerted on cyprid metamorphosis of B. amphitrite was examined. Extracts of algae and associated bacteria showed no effect on the metamorphosis of B. amphitrite. This may be attributed to absence of cue‐specific sugars in the exopolysaccharides and culture supernatants of bacteria. On the other hand, natural leachants of M. tintinnabulum, which showed the presence of N‐acetyl‐d ‐glucosamine, a known algal spore attractant, also inhibited metamorphosis of B. amphitrite. Thus, hosting specific epibionts could have important roles in the segregation of barnacle population.     

Macrofouling communities in the cooling system of the Vladivostok heat and power plant

The composition, structure, and distribution of flora and fauna on hydraulic facilities in the cooling system of the Vladivostok combined heat and power plant-2 were studied in summer and autumn of 2001. Cluster analysis was applied to differentiate Balanus rostratus, Mytilus trossulus, Jassa marmorata, Crassostrea gigas + Balanus rostratus, Modiolus modiolus + Pachycheles stevensii, Hydroides ezoensis, and Amphibalanus improvisus communities. The groups of fouling organisms found on the studied objects were shown to be assigned both to “physically controlled” and “biologically balanced” benthic communities. This study stresses the high similarity between the fouling communities of anthropogenic substrata and natural intertidal and subtidal benthic communities from adjacent areas of Peter the Great Bay.     

Seasonal variation of zinc in the barnacle, Balanus balanoides (L.) maintained on a raft in the Menai Strait

A population of the intertidal barnacle, Balanus balanoides (L.) maintained on a raft in the Menai Strait was sampled regularly over a two-year period and pooled fractions analysed for zinc and other trace metals. The seasonal variation of zinc in three fractions (body, other soft tissues and shell) has been followed and related to physiological condition. The body accumulates zinc almost continuously whilst this trace metal accumulates in the ovary only as it develops and is lost in the gametes when these are laid. The newly formed egg-masses therefore may be the more useful sample for indicating zinc pollution.     

The Southwest Atlantic intertidal burrowing crab Neohelice granulata modifies nutrient loads of phreatic waters entering coastal area

Along the coastal areas of the Southwest Atlantic estuaries and embayments, phreatic water often circulates through very extended areas (up to several hundred meters perpendicular to the coast), dominated by dense assemblages of deep burrows of the crab Neohelice granulata (formerly Chasmagnathus granulatus). This crab inhabits the intertidal area, from mudflats to marshes vegetated by species of Spartina, Sarcocornia and Juncus, generating extensive burrowing beds where burrow density may reach up to 60 burrows m⁻². Since the lower limit of the crab burrows is usually the water table, we investigated through field experiments the effect of N. granulata and their burrows on the chemical characteristics of this phreatic water. Water analysis from experimental (1) occupied burrows (with crabs), (2) unoccupied burrows (where crabs were excluded), and (3) sediment pore water show remarkable differences. Water oxygenation, and nitrate, ammonium and sulphate concentrations inside occupied burrows were higher than in the water inside unoccupied burrows or pore waters. Moreover, directed sampling of phreatic water entering and leaving the crab bed, shows that dissolved inorganic nitrogen concentration is enhanced as the water crosses the crab bed. These results may be ascribed to the fact that in the salt marsh the crabs spend most of their time within burrows, where presumably they store food (plants) and defecate. These activities generate an area of accumulation of excrements and nutrients in different decomposition states. The present work shows a novel way by which bioturbating organisms can affect nutrients exportation from salt marshes to the open waters.     

Predatory impact of an epiphytic hydrozoan in an upwelling area in the Bay of Coliumo (Dichato, Chile)

The natural diet of the epiphytic hydrozoan Obelia geniculata has been studied in an upwelling area in the Bay of Coliumo (Dichato, Chile) during two 24-h cycles. Number of prey per predator and predatory density have been measured. In both cycles more than 78% of the diet consisted of eggs of invertebrates and faecal pellets. Compared with other species of hydrozoans studied in a similar way, O. geniculata had a heterogeneous diet and a capture rate which was scarcely related to the peaks of abundance of its prey. This catch rate was between 632×10³ and 10 393×10³ prey m⁻² day⁻¹, which corresponds to a mean ingestion rate of 113% of the hydranth biomass per day. These results indicate the importance of small-sized benthic suspension feeders in upwelling systems.     

The trophic position of the alien crab <Emphasis Type="Italic">Rhithropanopeus harrisii</Emphasis> (crustacea decapoda panopeidae) in the Taman Bay,Sea of Azov community

This work concerns the trophic web positioning of the alien crab Rhithropanopeus harrisii and other common marine invertebrate species and fishes in the benthic ecosystem of the shallows of Taman Bay, Sea of Azov. The base of the trophic web in this system is composed of phytoplankton, macrophytes (algae and marine grasses), and reeds that use atmospheric carbon for photosynthesis. Analysis of the isotopic composition of nitrogen and carbon has shown that although marine grasses are dominating primary producers in the shallows of the bay, primary consumers (such as Cerastoderma glaucum, Porifera gen. sp., Gammarus aequicauda, Deshayesorchestia deshayesii and Idotea balthica) only partially use this organic source; instead, they use a combination of different sources of primary production. It has been shown that the food source of the alien crab is primarily of animal origin. In Taman Bay, R. harrisii is on the same trophic level as other carnivores/scavengers: benthic fishes Syngnathus nigrolineatus, Gobius spp. and native crab Pilumnus hirtellus and shrimp Palaemon adspersus.     

Trophic relationships in deep-water decapods of Le Danois bank (Cantabrian Sea,NE Atlantic): Trends related with depth and seasonal changes in food quality and availability

The trophic relationships of decapod crustaceans on Le Danois bank (NE of Iberian Peninsula, NE Atlantic Ocean) were studied within the framework of the multidisciplinary project ECOMARG during two surveys, one in October 2003 and the other in April 2004. The diets of eleven species of decapods were analyzed and, within a rather continuous gradient of food source exploitation, 3 trophic groups were identified: (1) plankton feeders, comprising the shrimps Acanthephyra pelagica, Sergia robusta, and Pasiphaea tarda, which preyed on meso-bathypelagic taxa such as euphausiids and calanoids; (2) benthos feeders, comprising the crangonids Pontophilus norvegicus and Pontophilus spinosus, the crab Geryon trispinosus and the shrimp Aristeus antennatus; and (3) an intermediate group, including the rest of species, with mixed diets that included detritus. Among the third group, anomurans (Munida tenuimana, Pagurus alatus, and Parapagurus pilosimanus) consumed phytoplanktonic detritus in April, suggesting a link with peaks of surface Chl a occurring between March and April in the study area. Gut pigment and isotopic (δ¹³C/δ¹⁵N correlations) analyses revealed that assemblages inhabiting the top of the bank (455–612 m) and the inner basin (642–1048 m, close to the Lastres canyon head) had different food sources, with species inhabiting the deepest region exhibiting a stronger dependence on marine snow derivatives. These results are consistent with the higher proportion of mud and sediment organic matter (OM) content in the inner basin (82.2% pellites; 6.3% OM at 1028 m) compared to the top of the Le Danois bank (only 13.9% pellites; 2.8% OM at 485 m), which is a hydrodynamically more active zone. Exploitation of different food sources is also consistent with differences in the trophic level of species, inferred from stable δ¹⁵N isotope analyses, which yield values ranging from 6.88‰ for the hermit crab P. alatus to 13.52‰ for the crangonid shrimp P. norvegicus. Stomach fullness was higher in April 2004 than in October 2003, both between and within species of the dominant decapods, including detritus feeders (M. tenuimana) and benthos feeders (e.g. G. trispinosus, P. norvegicus). Most species exhibited a parallel increase in their density in April 2004, with a significant positive correlation between density and stomach fullness. This increase coincides with a peak of surface Chl a concentration occurring in March–April.     

Colonization process by macrobenthic infauna after a managed coastal realignment in the Bidasoa estuary (Bay of Biscay, NE Atlantic)

The colonization process by macrobenthic infauna after the completion of managed realignment on the fertile plain of Jaitzubia, along the Bidasoa estuary, is assessed here. The benthic invertebrate fauna was sampled annually for three years in the newly recovered intertidal areas. The results show a rapid colonization process. Six months after the completion of the restoration works, all the stations exhibited a large number of small opportunistic benthic species, as well as some large individuals of Hediste diversicolor, that dominated the biomass of the community. One year later, a new stage of development or succession towards the settlement of the Scrobicularia plana–Cerastoderma edule community was observed, showing a reduction of both abundance and species richness, and an increase in biomass, mainly due to the presence of individuals of S. plana (which were absent previously). The following year, a further reduction in the abundance and number of species was observed, as well as an increase in biomass, but on this occasion S. plana was the main contributor to community biomass. Evidence of the importance of passive or active dispersion in the adult stage of the species H. diversicolor and S. plana in the recovery process was also observed.     

Long-term changes in the production by estuarine macrobenthos affected by multiple stressors

The macrobenthic production of an estuarine system was evaluated over a 14-year study period in a seagrass bed and in a sandflat. Over this period, the estuary suffered severe eutrophication and extreme weather events with important impacts on the community, impairing system functioning and ultimately the goods and services provided by the estuary (decline in the seagrass bed, decreased community production and/or a boost in the production by opportunist species, such as Hydrobia ulvae). Following the anthropogenic impacts, management measures were introduced which allowed a gradual recovery of the seagrass bed and a new macrobenthic community structure manifested by production increases of slow-growing species, such as Scrobicularia plana and Hediste diversicolor. There was a gradual re-orientation of energy into population biomass instead of population density but this was not translated into higher community production, mainly due to the decreased production of opportunist species (H. ulvae and several polychaetes). Several weather extreme events occurred during this post-mitigation phase - floods, heatwaves and droughts, all of which had negative impacts on macrobenthic dynamics and production. The heatwaves led to the greatest decreases in macrobenthic production, mainly due to S. plana perhaps associated with its physiological intolerance of higher temperatures. The prolonged drought that followed the heatwaves maintained low levels of production by S. plana and H. ulvae. With climate change, the frequency and intensity of extreme weather events are likely to increase worldwide so that the recovery of impacted/disturbed systems from impacts such as eutrophication may be seriously affected by these additional stressors, compromising attempts to improve the ecological quality of estuarine ecosystems.     

A rapid assessment survey of invasive species of macrobenthic invertebrates in Korean waters

Introduced species are a growing and imminent threat to living marine resources in parts of the world’s oceans. The present study is a rapid assessment survey of invasive macrobenthic invertebrate species in Korean ports. We surveyed over 40 ports around Korea during the period of May 2010~March 2013. Among the sampling sites were concrete walls, docks and associated floats, bumpers, tires, and ropes which might harbor non-native species. We found 15 invasive species as follows: one Sponge, two Bryozoans, three Mollusks, one Polychaete, four Cirripedes, and four Ascidians. Three morphologically similar species, namely X. atrata, M. galloprovincialis, and X. securis were further examined for distinctions in their morphology. Although they could be reasonably distinguished based on shell shapes, significant overlap was noted so that additional analysis may be required to correctly distinguish them. Although many of the introduced species have already spread to all three coastal areas, newly arrived invasive species showed a relatively restricted range, with a serpulid polychaete Ficopomatus enigmaticus and a mytilid bivalve Xenostrobus securis found only at a few sites on the East Coast. An exception is for Balanus perforatus, which has rapidly colonized the East coast of Korea following its introduction into the region. Successful management of invasive macrobenthic invertebrates should be established in order to contain the spread of these newly arrived species.     

Gastropod shell repair tracks predator abundance

Predator‐prey interactions can shape biological communities and drive evolutionary change. The assessment of predation rates in modern and ancient ecosystems presents many challenges. We examined the utility of one common predation metric, shell repair frequency, as a proxy for crushing predation on marine gastropod populations. Our six localities near Bamfield, British Columbia, represented two contrasting predation regimes: three high‐wave‐energy (exposed) environments have low predation rates, while three low‐wave‐energy (protected) environments exhibit high predation rates. We measured the occurrence of repair scars for local populations of the gastropod Chlorostoma funebrale, and recorded multiple independent variables (shell measurements, water velocity, predatory crab abundance) at each locality. Fisher's exact test established that the repair frequencies observed in the protected localities were significantly greater than those of the exposed environments. Factors important in predicting the likelihood of an individual having a repair scar were examined using binomial logistic regression. A model containing the main effects of crab abundance, shell height, shell thickness and water velocity provided the best fit in predicting the presence of repair scars, with crab abundance having the largest contribution. The strong relationship between the presence of repair scars and predator abundance indicates that repair frequency is a valid tool for assessing predation intensity in gastropod populations.     

Seasonal dynamics of the density of the crab larvae (Decapoda: Brachyura et Anomura) in Minonosok Bay of Pos’eta Bay (Peter the Great Bay, Sea of Japan)

As the materials for this project, we used data on the distribution of the larvae of the Asian paddle crab Charybdis japonicus (A. Milne-Edwards, 1861); the spider crabs Hyas coarctatus ursinus (=Hyas ursinus) (Rathbun, 1924), Pugettia quadridens (de Haan, 1839), and Pisoides bidentatus (H. Milne-Edwards, 1873); the samurai crab Paradorippe granulata (=Dorippe granulata) (de Haan, 1841); the pea crab Pinnixa rathbuni (Sakai, 1934); and the porcelain crab Pachycheles stevensii (Stimpson, 1858) in Minonosok Bay of Pos’eta Bay obtained during 2000–2002 and in 2004. The planktonic samples were collected from the last third of May to September. The greatest density of the larvae was observed in May, mid-June, late June-early July, and late July-early August. The greatest densities of the crab larvae and the decapod larvae ranged from 20.4 to 48.2 and from 88.4 to 245.3 specimens/m³. The schedule of the crab larvae’s occurrence in the plankton is provided for the first time. The distribution of the density showed pronounced patchiness.