Patterns of larval release in the Florida stone crab, Menippe mercenaria

Larval release patterns in brachyuran crabs are often synchronized with environmental cycles. While previous studies have focused extensively on supratidal and intertidal taxa, there have been relatively few investigations of subtidal species. This study examined patterns of larval release by the Florida stone crab, Menippe mercenaria, from three different tidal regimes. Ovigerous stone crabs were collected from Sebastian Inlet on the east coast of Florida, Tampa Bay on the west coast of Florida, and the Florida Keys. Patterns of larval release were monitored in the laboratory in relation to local tidal and diel cycles. Results showed a significant diel pattern in initiation of hatching by crabs from each of three study areas. Larval release consistently occurred during the diurnal phase despite the maintenance of females in constant laboratory conditions for up to 96 h prior to hatching. This implies that release may be controlled by a circadian clock. Patterns of release by stone crabs in relation to tidal cycle were more variable. Larval release by females from populations near Tampa Bay and Sebastian Inlet were not synchronized with the tides, whereas females collected from the Florida Keys exhibited a pattern that was strongly related to tidal cycle. These results may be explained by differences in tidal amplitude at the three sampling locations.     

The influence of plankton composition and water quality on hard clam (Mercenaria mercenaria L.) populations across Long Island's south shore lagoon estuaries (New York, USA)

We conducted a two-year study to assess how plankton composition and water quality impacts the distribution, densities, condition, growth, biochemical composition and reproductive success of juvenile and adult Mercenaria mercenaria (L.) in Long Island's south shore estuaries (LISSE). Juvenile and adult hard clams were placed in suspended cages at 10 locations ranging from the ocean inlets to locations furthest from inlets in Shinnecock Bay (SB), the eastern-most barrier island estuary of LISSE, and Great South Bay (GSB), the western-most barrier island estuary of LISSE. Phytoplankton community composition, temperature, salinity, dissolved oxygen, and clam growth and condition were monitored bi-weekly. A benthic survey of M. mercenaria densities in both estuaries was also conducted. In both 2004 and 2005, juveniles in central bay locations had significantly faster growth rates, lower mortality rates, and higher lipid content relative to sites closest to the inlets. Adult hard clams closest to the Fire Island inlet also had significantly lower condition indexes compared to mid-bay stations and densities of wild M. mercenaria populations in both estuaries were lower near inlets compared to locations further from inlets. In addition to substantial spatial differences within each estuary, differences were also observed between the embayments as juvenile clams in SB grew approximately twice as fast as those in GSB and adults in SB had significantly greater condition indexes than clams in GSB. Instantaneous juvenile growth rates were highly correlated to temperatures below 24 °C (p < 0.0001) and were also significantly correlated with several indicators of suspended food quantity and food quality (centric diatoms, phytoplankton cells > 5 μm, and dinoflagellates (inverse correlation)) which co-varied independently of temperature. In sum, these results suggest tidal exchange in LISSE promotes a water quality regime (cold water, with low food concentration) which would reduce the growth of juvenile clams and the overall reproductive success of adult hard clams located near newly-formed ocean inlets. However, increased exchange for regions furthest from inlets could enhance juvenile clam growth rates by reducing summer peak temperatures (> 24 °C) and densities of poor food sources (dinoflagellates).     

Multiple cues from multiple habitats: Effect on metamorphosis of the Florida stone crab, Menippe mercenaria

The Florida stone crab, Menippe mercenaria, is an economically and ecologically important species that ranges from North Carolina throughout the Caribbean and the southeastern Gulf of Mexico. However, there is little known about its early life history stages as compared to other commercially important species in the region. The goal of this research was to examine effects of putative cues on metamorphosis from the megalopa stage to the first juvenile stage. Our study investigated the effect of water-soluble exudates from four substrata, as well as natural biofilms, and exudates from adult stone crabs. In addition, the influence of natural substrata was compared to that of artificial substrata. Adult exudate had no significant effect on metamorphosis, despite a wide range of tested concentrations. In contrast, there was a significant effect on mean time to metamorphosis in experimental groups exposed to multiple cues associated with the brown alga Sargassum fluitans, rubble from stone crab habitat, the eastern oyster Crassostrea virginica, and biofilms associated with the oyster. Furthermore, we provide evidence for metamorphic responses to water-soluble chemical cues, as well as biochemical and physical cues associated with different substrata. Overall results were coherent with the relevant body of previous work on metamorphosis of brachyuran crab larvae and indicate that both physical and chemical cues are important factors in facilitating the settlement and metamorphosis of M. mercenaria larvae in juvenile nursery habitat.     

Potential impacts of brown tide, Aureococcus anophagefferens, on juvenile hard clams, Mercenaria mercenaria, in the Coastal Bays of Maryland, USA

The rate of growth of juvenile hard clams, Mercenaria mercenaria, was studied in the Coastal Bays of Maryland during an outbreak of the brown tide, Aureococcus anophagefferens. Brown tide dominated the plankton community during the month of June 2002, with cell densities at several sites reaching category 3 (>200,000 cells ml⁻¹) levels. Temperatures during the bloom were 18.6–27.5 °C. Nutrient conditions preceding and during the bloom were conducive for the proliferation of A. anophagefferens: while inorganic nitrogen and phosphorus were <1 μg at N or P l⁻¹, urea was elevated during bloom development. Organic nitrogen, phosphorus and carbon were in the range of levels observed in previous brown tide blooms and increased following the collapse of the bloom. Growth rates of juvenile clams were significantly lower during the period of the brown tide bloom than following its collapse. Growth rates of M. mercenaria were found to be negatively impacted at brown tide densities as low as 20,000 cells ml⁻¹, or category 1 levels. The low growth rates of M. mercenaria could not be explained by temperature, as the lowest growth rates were found when water temperatures were at levels previously found to be optimal for growth.     

Chemically induced predator avoidance behaviour in the burrowing bivalve Macoma balthica

The responses of the burrowing bivalves Macoma balthica and Cerastoderma edule to chemical cues emitted by feeding shore crabs Carcinus maenas were investigated. M. balthica held in the laboratory and exposed to chemical signals in effluent water discharging from tanks containing C. maenas fed 20 M. balthica day^− 1 reacted by increasing their burial depths from approximately 30 mm to depths of > 60 mm, over a period of several days. When the signal was removed the bivalves gradually returned to their original depth over 5 days. C. edule similarly exposed to effluent from crabs feeding on conspecifics showed no response. In an attempt to identify the signal inducing this burrowing response, M. balthica were exposed to a variety of chemical signals. Crabs fed M. balthica elicited the strongest response, followed by crabs fed C. edule. There were also small responses to effluent from crabs fed on fish, crabs previously fed on M. balthica and to crab faeces, but no responses to starved crabs, crushed M. balthica, or controls. We conclude that increased burrowing depth of M. balthica is induced by some as yet unidentified chemical cue produced by feeding crabs and is strongest when the crabs were fed on M. balthica. Unexpectedly, neither the presence of crabs themselves, nor of damaged conspecifics, was effective in eliciting a burrowing response. The mortality rates of M. balthica and C. edule selected by crabs when burrowed at normal depths and after exposure to effluent from feeding crabs were different. Crabs selected 1.5 times more C. edule than M. balthica when both species were burrowed at their normal depths, but 15 times more after the tanks had been exposed to effluent from feeding crabs for 5 days. The burrowing response of M. balthica thus appears to reduce mortality significantly by displacing predation pressure on to the more accessible C. edule.     

Sex-specific differences and the role of predation in the interaction between the hermit crab, Pagurus longicarpus, and its epibiont, Hydractinia symbiolongicarpus

The interaction between the hermit crab, Pagurus longicarpus, and the shell epibiont, Hydractinia symbiolongicarpus, varies from mutualism to parasitism based on the environmental context. We tested the hypothesis that this interaction also varies as a function of hermit crab sex. Given that recent work showed a negative effect of Hydractinia on female reproduction, we predicted a greater frequency of males in Hydractinia shells in the field and a stronger preference by males than females for shells with Hydractinia. Field collections documented a significantly greater proportion of males than females occupying shells with Hydractinia, and a significantly greater proportion of ovigerous females than non-ovigerous females in shells with Hydractinia. In laboratory shell-switching experiments, a greater proportion of males than females chose to enter shells with Hydractinia, but there was no difference in the proportions of males and females that vacated shells with Hydractinia.We examined whether the presence of Hydractinia influenced predation rates. Blue crabs fed on more than twice as many hermit crabs in shells with Hydractinia as compared to bare shells, but there was no significant difference for stone crabs. Laboratory experiments showed that the force required to crush shells was significantly greater for shells without Hydractinia. Thus, the lower occupancy and preference exhibited by females than males for shells with Hydractinia appears to result both from the decreased reproduction shown in past studies and an increase in predation risk.     

Reduction of attractiveness to the sea star Asterias forbesi (Desor) by the clam Mercenaria mercenaria (Linnaeus)

Both in field and laboratory choice tests, the sea star, Asterias forbesi (Desor), was attracted to distant upstream clams, Mercenaria mercenaria (Linnaeus). Clams exposed to upstream sea stars were chosen less frequently by downstream sea stars than clams without sea stars upstream. Sea stars neither attracted nor repelled downstream conspecifics.When clams were exposed to upstream sea stars, their oxygen consumption decreased, as did their pumping rate and activity (as measured by number of visible siphons). The former may result from one or both of the latter.It is concluded that clam and sea star sense each other over a distance by chemical cues. The response of the clam is a general lowering of activity which may result in decreased attractiveness to sea star predators. This response may serve as a defensive measure against distance detection by Asterias forbesi.     

Effects of temperature on hard clam (Mercenaria mercenaria) immunity and QPX (Quahog Parasite Unknown) disease development: II. Defense parameters

Quahog Parasite Unknown (QPX) is a protistan parasite affecting hard clams Mercenaria mercenaria along the Northeastern coast of the United States. The geographic distribution and occurrence of disease epizootics suggests a primary role of temperature in disease development. This study was designed to investigate the effect of temperature on constitutive and QPX-induced defense factors in M. mercenaria. Control and QPX-challenged (both experimentally and naturally) clams were maintained at 13, 21 and 27 °C for 4 months. Control and experimentally-infected clams originated from a southern broodstock (Florida, no prior reports of disease outbreak) while naturally-infected clams originated from a northern broodstock (Massachusetts, enzootic area). Standard and QPX-specific cellular and humoral defense parameters were assessed after 2 and 4 months. Measured parameters included total and differential hemocyte counts, reactive oxygen species production, phagocytic activity of hemocytes, lysozyme concentration in plasma, anti-QPX activity in plasma and resistance of hemocytes to cytotoxic QPX extracellular products. Results demonstrated a strong influence of temperature on constitutive clam defense factors with significant modulation of cellular and humoral parameters of control clams maintained at 13 °C compared to 21 and 27 °C. Similarly, clam response to QPX challenge was also affected by temperature. Challenged clams exhibited no difference from controls at 27 °C whereas different responses were observed at 21 °C and 13 °C compared to controls. Despite differences in infection mode (experimentally or naturally infected) and clam origin (northern and southern broodstocks), similarities were observed at 13 °C and 21 °C between QPX infected clams from Florida and Massachusetts. Clam response to temperature and to QPX exhibited interesting relationship with QPX disease development highlighting major influence of temperature on disease development.     

The SW Atlantic burrowing crab Chasmagnathus granulatus Dana affects the distribution and survival of the fiddler crab Uca uruguayensis Nobili

In this paper, we address the question of whether the presence of the burrowing crab Chasmagnathus granulatus affects the habitat use of the fiddler crab Uca uruguayensis. Field samples showed that the species have a disjoint spatial distribution. Male fiddler crab density decreased in zones with C. granulatus, however, female density increased. Male fiddler crabs avoided feeding on sediment affected by C. granulatus and were more preyed. Predation was higher during the fiddler crab reproductive season and, probably due to predation risk, males showed lower reproductive display in shared zones. Field experiments shows that when C. granulatus were excluded, densities of U. uruguayensis increased mainly due to an increase in density of males. Habitat differentiation of these species may be because C. granulatus affects U. uruguayensis in several ways, including direct predation, disturbance and behavioural changes associated to predation risk. Males and females are affected differentially probably because of the extreme sexual dimorphism of this crab species. Coloration on enlarged claw and waving activities are all factors that increase predation risk for male and the presence of only one feeding claw may increase sediment-mediated effects.     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Interaction strength between a predator and dangerous prey: Sinistrofulgur predation on Mercenaria

The lack of direct empirical evidence of predator evolution in response to prey adaptation is a fundamental weakness of the arms race analogy of predator-prey coevolution. I examined the interaction between the predatory busyconine whelk Sinistrofulgur sinistrum and its bivalve prey Mercenaria mercenaria to evaluate whether reciprocal adaptation was likely in this predator-prey system. Thick-lipped whelks use their shell lip to chip open the shell of their prey, often resulting in breakage to their own shell. Thus, hard-shelled prey, such as Mercenaria, may be considered dangerous because they are able to inflict damage to the predator as a consequence of the interaction. The strength of interaction between whelks and their bivalve prey was viewed by regressing predator performance (the incidence of shell breakage in encounters with prey) on prey phenotype (a function of size). Interaction with Mercenaria of varying sizes has strong and predictable consequences (r²=0.946; p=0.028) for Sinistrofulgur. Predators that select large, thick bivalve prey increase the likelihood that their shell lip will be broken in the process of attempting to open their prey. Ecological consequences of feeding-induced breakage may include reduced growth rate, reproductive success, and survivorship. These results suggest that natural selection should favor predator phenotypes that reduce feeding-induced breakage when interactions with damage-inducing prey occur.     

Distances of dispersal of juvenile bivalves (Mya arenaria (Linnaeus), Mercenaria mercenaria (Linnaeus), Gemma gemma (Totten))

Although it is recognized that many species of benthic invertebrates continue to disperse after settlement, particularly in soft-bottom habitats, the scale over which movements of juveniles occur is not well known. This study combined laboratory flume experiments assessing the effects of clam size, species, and water velocity on rates and distances of dispersal of three species of juvenile bivalves with field measurements of loss rates and distances of dispersal of transplanted bivalves in the Navesink River estuary in New Jersey, USA. Dispersal distances measured in the laboratory ranged from an average of 1.6 to 40 cm h^− 1 depending on clam size, species, and flow speed. Distances and likelihood of dispersal were generally greater for Mya arenaria than for Mercenaria mercenaria or Gemma gemma, although differences between species were not consistent. As predicted, smaller (1.3 mm) M. arenaria tended to disperse more than larger (3.7 mm) ones, although no significant differences were detected between two sizes (1.8 and 3.4 mm) of M. mercenaria. The similarity of the erosion thresholds of dead clams across sizes and species suggests that burrowing behaviour plays an important role in determining variation in dispersal due to clam size and species. In the field, densities of clams (M.arenaria and M.mercenaria) were reduced to half of that in controls after 3.5-5 h, indicating high levels of dispersal and/or mortality. Some individuals were recovered up to 50 cm away from their initial locations. Overall, our results suggest that dispersal distances of these three species due to bedload transport are likely to be on the order of centimeters per hour. Although these dispersal distances are small, such movements are likely to occur frequently due to tidal currents and, consequently, may have profound impacts on patterns of abundance and distribution.     

Effects of substrate on interactions between juvenile sea scallops (Placopecten magellanicus Gmelin) and predatory sea stars (Asterias vulgaris Verrill) and rock crabs (Cancer irroratus Say)

We investigated the effect of substrate (glass bottom, sand, granule, pebble) on predation of juvenile sea scallops (Placopecten magellanicus) by sea stars (Asterias vulgaris) and rock crabs (Cancer irroratus) at two prey sizes (11-15 mm and 24-28 mm shell height), and two prey densities (10 and 30 scallops per aquarium) in laboratory experiments. Specifically, we quantified predation rate and underlying behaviours (proportion of time a predator spent searching for and handling prey, encounter rate between predators and prey, and various outcomes of encounters). We detected a significant gradual effect of particle size of natural substrates on sea star predation: specifically, predation rate on and encounter rate with small scallops tended to decrease with increasing particle size (being highest for sand, intermediate for granule, and lowest for pebble). Substrate type did not significantly affect predation rates or behaviours of sea stars preying on large scallops or of rock crabs preying on either scallop size classes. Other factors, such as prey size and density, were important in the scallop-sea star and scallop-rock crab systems. For example, predation rate by sea stars and crabs and certain sea star behaviours (e.g. probability of consuming scallops upon capture) were significantly higher with small scallops than with large scallops. As well, in interactions between small scallops and sea stars, predation rate and encounter rate increased with prey density, and the proportion of time sea stars spent searching was higher at low prey density than high prey density. Thus, substrate type may be a minor factor determining predation risk of seeded scallops during enhancement operations; prey size and prey density may play a more important role. However, substrate type still needs to be considered when choosing a site for scallop enhancement, as it may affect other scallop behaviours (such as movement).     

Large- and small-scale effects of habitat structure on rates of predation: how percent coverage of seagrass affects rates of predation and siphon nipping on an infaunal bivalve

Landscape ecology, predominantly a terrestrial discipline, considers the effect of large-scale (tens of meters to kilometers) spatial patterns of habitats on ecological processes such as competition, predation, and flow of energy. In this study, a landscape-ecology approach was applied to a marine soft-sediment environment to examine rates of predation and transfer of secondary production in and around vegetated habitats. Seagrass beds naturally occur in a variety of spatial configurations from patches 1–10s of meters across with interspersed unvegetated sediments (i.e., patchy coverage) to more continuous coverage with little or no bare sediment. I designed experiments to address how percent coverage of seagrass in a 100-m² area of seafloor, and the spatial arrangement (degree of patchiness or fragmentation) of an equal area (100 m²) of vegetation affected predation (lethal) and siphon nipping (sublethal) intensity on an infaunal bivalve, Mercenaria mercenaria (hard clam). Measures of seagrass density and biomass with different percent coverage of seagrass were also made. When clams were placed in both the vegetated and unvegetated portions of the seafloor nearly twice as many clams were recovered live with 99% seagrass cover than with 23% seagrass cover, while survivorship was intermediate with 70% cover. Cropping of clam siphons from both the vegetated and unvegetated sediments was also affected by the amount of seagrass cover in a 100-m² area of seafloor: mean adjusted siphon weights were approximately 76% heavier from the 99% seagrass cover treatment than from the 70% or 23% cover treatments. Survivorship of clams placed within an equal area of seagrass in very patchy, patchy, and continuous spatial configurations was 40% higher in the continuous seagrass treatment than in either of the two patchy treatments. This study demonstrates that transfer of secondary production in the form of predation and cropping on an infaunal organism is altered as the percent cover of seagrass changes. While large-scale changes in the amount and spatial patterning of vegetation may affect habitat utilization patterns and foraging HGLoopbehavior, increased seagrass density and biomass with increased percent coverage of seagrass limit any conclusions concerning predator foraging behavior and feeding success in response to patch shapes and sizes. Instead, local changes in seagrass characteristics provide the most compelling explanation for the observed results.     

Changes in shell and soft tissue growth, tissue composition, and survival of quahogs, Mercenaria mercenaria, and softshell clams, Mya arenaria, in response to eutrophic-driven changes in food supply and habitat

Eutrophic-driven changes in the composition of near-bottom seston and surface sediment potentially affect food resources and habitat of commercially important bivalves like quahogs, Mercenaria mercenaria, and softshell clams, Mya arenaria. To define how land-derived nitrogen loads and resulting eutrophication affect bivalves, we compared estuarine features to growth and survival of clams across estuaries receiving different N loads. The major effects of nitrogen enrichment on near-bottom seston and surface sediment were to (1) increase microalgal concentrations and reduce carbon to nitrogen ratios, increasing quantity and quality of available foods, and (2) reduce oxygen content in sediments, potentially reducing habitat quality. Shell growth of juvenile and native clams increased with increasing food supply, driven by N enrichment. Growth of soft tissue followed growth of shell, and %N content of soft tissue increased across N loads, providing direct evidence of a link between N loads and growth responses in clams. In some locations, low salinity limited growth and low oxygen concentrations may have reduced survival. Despite these factors, our data indicate the major effect of N enrichment on clams was increased secondary production in terms of shell and soft tissue growth.     

Predator-prey interaction between hatchery-reared Japanese flounder juvenile, Paralichthysolivaceus, and sandy shore crab, Matutalunaris: daily rhythms, anti-predator conditioning and starvation

Predator-prey interaction between sandy shore crab, Matuta lunaris (Forskål, 1775), and juvenile Japanese flounder, Paralichthys olivaceus (Temminck et Schlegel), was investigated under controlled laboratory conditions. Possibility of training and conditioning hatchery-reared flounder to avoid predators was also examined. Crabs took over 75% of their daily ration at night when they were given access to prey 24 h a day. Large (64.8±5.4 g)- and medium (30.68±3.33 g)-sized crabs ate ca. 5.5±1.45 and 3.9±1.99 individuals of flounder (TL=4.96±0.23 cm) a day, respectively. When flounder juveniles that have experienced predation pressure by crabs encountered predators again, they exhibited better survival compared to the naive fish. Flounder juveniles were also conditioned either using small and, thus, benign predators, or large crabs over fence. The conditioned fish with either method were better able to avoid capture by crabs than naive fish, revealing that learning process should play an important role in their predator avoidance. Anti-predator performance was also compared between starved and fed flounder juveniles. Fed fish were rarely eaten by predators after 3 h of exposure, whereas starved fish continued to be eaten. Our results suggest that stock-enhancement program of Japanese flounder can be improved by applying proper feeding protocol and conditioning to avoid predators prior to release. Present research supports the idea that behavioural and ecological consideration for the target species is indispensable for the success of stock enhancement.     

Modification of animal habitat by large plants: mechanisms by which seagrasses influence clam growth

Summary Field experiments withMercenaria mercenaria in a relatively high-energy environment demonstrated that clams on unvegetated sand flats failed to grow during autumn while those within seagrass beds grew substantially. Clam growth rates at the seagrass margin that first receives the faster-flowing, flood-tidal currents were about 25% less than at the opposite edge. In a second experiment, pruning, which reduced average blade length by 50–75%, was shown to enhance near-bottom current velocities and to reduce shell growth ofMercenaria during summer by about 50%. As in the first experiment, clams in the unvegetated sand flats exhibited no net growth. Clam mortality, caused mostly by predatory crabs and whelks, was much higher on sand flats than in seagrass beds and intermediate in clipped seagrass. Although consistent with some previous reports, these growth results are still surprising given that they contradict the generalization that suspension feeders grow faster under more rapid current regimes.Three types of indirect interactions might explain the observed effect of seagrass on growth of buried clams: (1) altering food supply; (2) changing the intensity of biological disturbance on feeding clams; and/or (3) affecting the physical stability of the sediments. Previous research on this question has focused almost exclusively on processes that alter food supply rates. In this study, food concentrations, as indicated by suspended chla, were 30% higher inside than outside one seagrass bed, whereas chla concentrations in two other beds were not different from those on adjacent sand flats. This result is sufficient to show that more intense food depletion was not induced by the reduction in flow velocities under the seagrass canopy. Nevertheless, the possible small difference in food concentrations between vegetated and unvegetated bottom seems insufficient to explain the absence of growth of sand-flat clams, especially given the virtual lack of food limitation among suspension feeders in this system. Two data sets demonstrated that the effects of biological disturbance agents cannot be ignored. An outdoor laboratory experiment showed that even in the absence of physical contact between predator and prey the presence of a whelk reduces the amount of time spent feeding byMercenaria. This result suggests that sand flats, where predation rates are higher, may be sites of lower clam growth than seagrass beds because of greater consumer interference with clam feeding. Furthermore, clam siphons are proportionately larger inside seagrass than on sand flats, implying that siphon nipping may not be as intense inside seagrass. This process, too, would reduce net growth of sand-flat clams. Finally, no explicit test was conducted of the hypothesis that enhanced sediment transport in the absence of flow baffling and root binding by seagrass inhibits net growth of clams on high-energy sand flats. Nevertheless, this is a reasonable explanation for the pattern of enhanced growth of seagrass clams, and could serve to explain the otherwise unexplained pattern of lower clam growth at the edge of the seagrass bed that experiences the faster flood-tidal current velocities. Each broad process, changing fluid dynamics, altering consumer access, and varying sediment stability, represents a mechanism whereby habitat structure, provided by the dominant plant, has an important indirect influence on the functional value of the habitat for resident animals.     

Effects of temperature on hard clam (Mercenaria mercenaria) immunity and QPX (Quahog Parasite Unknown) disease development: I. Dynamics of QPX disease

Quahog Parasite Unknown (QPX) causes disease and mortality in hard clams, Mercenaria mercenaria. Seasonality of QPX disease prevalence in the field and changes in QPX growth and survival in vitro suggest a role of temperature in the hard clam-QPX interaction and disease development. This study specifically examined the effect of temperature on QPX disease development and dynamics. Naturally and experimentally infected clams were separately maintained in the laboratory at 13 °C, 21 °C, or 27 °C for 4 months. Following this initial treatment, temperature was adjusted to 21 °C for 5 additional months to simulate seasonal changes of temperature in the field and to investigate the effect of temperature variations on QPX disease dynamics. Mortality was continuously monitored during the experiment and clams were sampled at 2, 4 and 9 months for the assessment of QPX disease prevalence and intensity using our standard histological and quantitative PCR techniques. Results demonstrated significantly higher QPX disease prevalence and intensity, as well as higher mortality, in naturally-infected clams maintained at 13 °C as compared to those held at 21 °C or 27 °C. Similarly, disease development was significantly higher in experimentally infected clams maintained at the colder temperature (70% prevalence after 4 months) as compared to those maintained under warmer conditions (<10%). Additionally, our results demonstrated an improvement in the condition of clams initially maintained at 13 °C for 4 months after transfer to 21 °C for 5 additional months, with a significant reduction of QPX prevalence (down to 19%). Interestingly, disease development or healing in clams maintained at different temperatures exhibited a strong relationship with clam defense status (jointly submitted paper) and highlighted the impact of temperature on clam activity and QPX disease dynamics. These findings should be taken into account for the timing of activities involving the monitoring, movement (e.g. relays, transplants) or grow out (e.g. commercial culture, municipal enhancement) of hard clams in enzootic areas.     

Hematodinium sp. infection of red Paralithodes camtschaticus and blue Paralithodes platypus king crabs from the Sea of Okhotsk, Russia

A disease caused by a parasitic dinoflagellate of the genus Hematodinium was identified in red, Paralithodes camtschaticus, and blue, Paralithodes platypus, king crabs from the north-east region of the Sea of Okhotsk, Russia, during annual stock surveys. No carapace color change was observed even in heavily infected crabs, but diseased crabs possessed creamy-yellow hemolymph, which was visible through the arthrodial membranes of the abdomen and appendages. Several stages of the parasite’s life history, including trophonts, plasmodia, sporonts and macrodinospores, were observed in tissues of infected king crabs. Numerous parasite cells were observed in the lumina of the myocardium, the gills, the connective tissue of antennal glands and the sinuses of nerve ganglia, eyestalks and gastrointestinal tract of king crabs with gross signs of infection. Based on sequencing of the 18S rDNA, it appears that the Hematodinium sp. found in red and blue king crabs is identical or closely related to Hematodinium sp. isolated from crabs of the genera Chionoecetes and Lithodes. Observed prevalences were 0.33% in sublegal male red king crabs, 0.18% in female red king crabs, 0.34% in sublegal male blue king crabs and 0.31% in female blue king crabs.