Factors influencing planktonic, post-settlement dispersal of early juvenile blue crabs (Callinectes sapidus Rathbun)

An emerging body of literature points to post-settlement, planktonic dispersal as a key determinant of distribution and abundance patterns of aquatic organisms, yet little is known about mechanisms inducing such dispersal. Recent evidence suggests that early juvenile blue crabs (Callinectes sapidus Rathbun) may use planktonic emigration as a means of post-settlement dispersal. The goal of this study was to identify mechanisms inducing post-settlement, planktonic dispersal of early juvenile blue crabs. A combination of field mark-recapture experiments in large seagrass beds within a 2x3 km region near Oregon Inlet, North Carolina, USA, and a series of laboratory flume experiments examined the effects of day vs. night, crab size (first-second juvenile benthic instars: J1-J2 vs. third-fifth juvenile benthic instars: J3-J5), crab density and current speed on planktonic dispersal of early juvenile blue crabs. Transport of dead crabs in the flume experiment identified that planktonic dispersal was an active behavioral response rather than a passive response to increasing current speed. The experimental results demonstrated that planktonic dispersal can range from 4 to 18% under medium to high flow conditions. Planktonic dispersal of juvenile crabs is (1) an active behavioral response, (2) increased significantly with current speeds above 20 cms(-1), and (3) was higher for relatively large (J3-J5) than small (J1-J2) instars. There was a non-significant trend towards greater dispersal at night than during the day in the field experiment, and no effect of crab density on dispersal in the flume experiment. The results from this study highlight the need to consider mechanisms inducing post-settlement, planktonic dispersal when attempting to understand and predict recruitment and population dynamics of aquatic organisms, as well as when linking hydrodynamics, animal behavior and planktonic dispersal.     

Endogenous swimming rhythms underlying secondary dispersal of early juvenile blue crabs, Callinectes sapidus

Blue crab, Callinectes sapidus Rathbun, megalopae settle in seagrass or other complex submerged aquatic habitats in estuaries, where they metamorphose to the first juvenile (J1) crab stage. Within tidal areas, early juveniles (J1-2) leave such nursery areas by undergoing secondary dispersal during nocturnal flood tides. The present study determined whether J1-2 blue crabs have a biological rhythm in vertical swimming activity that contributes to secondary dispersal. Endogenous rhythms in vertical swimming were determined for (1) J1-2 crabs collected from two estuaries with semi-diurnal tides, (2) J1 crabs that metamorphosed from the megalopal stage in the laboratory the day after collection, and (3) premolt megalopae that metamorphosed to J1 crabs under constant conditions during the experiment. In all cases, a circadian rhythm was present in which crabs swam vertically during the time of night in the field. The time of peak vertical swimming did not correspond to the time of flood tide at the collection sites, but did consistently occur at night, with a mean around midnight. While responses to environmental factors probably control the onset and end of vertical swimming by early juvenile blue crabs during flood tides in tidal areas, a circadian rhythm underlies secondary dispersal at night.     

Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance

Seagrasses form temporally dynamic, fragmented subtidal landscapes in which both large- and small-scale habitat structure may influence faunal survival and abundance. We compared the relative influences of seagrass (Zostera marina L.) habitat fragmentation (patch size and isolation) and structural complexity (shoot density) on juvenile blue crab (Callinectes sapidus Rathbun) survival and density in a Chesapeake Bay seagrass meadow. We tethered crabs to measure relative survival, suction sampled for crabs to measure density, and took seagrass cores to measure shoot density in patches spanning six orders of magnitude (ca. 0.25-30,000 m²) both before (June) and after (September) seasonally predictable decreases in seagrass structural complexity and increases in seagrass fragmentation. We also determined if juvenile blue crab density and seagrass shoot density varied between the edge and the interior of patches. In June, juvenile blue crab survival was not linearly related to seagrass patch size or to shoot density, but was significantly lower in patches separated by large expanses of unvegetated sediment (isolated patches) than in patches separated by <1 m of unvegetated sediment (connected patches). In September, crab survival was inversely correlated with seagrass shoot density. This inverse correlation was likely due to density-dependent predation by juvenile conspecifics (i.e. cannibalism); juvenile blue crab density increased with seagrass shoot density, was inversely correlated with crab survival, and was greater in September than in June. Shoot density effects on predator behavior and on conspecific density also likely caused crab survival to be lower in isolated patches than in connected patches in June. Isolated patches were either large (patch area >3000 m²) or very small (<1 m²). Large isolated patches had the lowest shoot densities, which may have allowed predators to easily find tethered crabs. Very small isolated patches had the highest shoot densities and consequently a high abundance of predators (=juvenile conspecifics). Though shoot density did not differ between the edge and the interior of patches, crabs were more abundant in the interior of patches than at the edge. These results indicate that seagrass fragmentation does not have an overriding influence on juvenile blue crab survival and density, and that crab cannibalism and seasonal changes in landscape structure may influence relationships between crab survival and seagrass habitat structure. Habitat fragmentation, structural complexity, faunal density, and time all must be incorporated into future studies on faunal survival in seagrass landscapes.     

The relative importance of habitat-specific settlement, predation and juvenile dispersal for distribution and abundance of young juvenile shore crabs Carcinus maenas L.

Young juveniles of many motile benthic species are concentrated in structurally complex habitats, but the proximate causes of this distribution are usually not clear. In the present study, I assessed three potentially important processes affecting distribution and abundance of early benthic stages in the shore crab (Carcinus maenas): (1) selection of habitat by megalopae (postlarvae); (2) habitat-specific predation; and (3) post-settlement movements by juveniles. These processes were assessed concurrently over 3-9 days at two spatial scales: at the scale of square meters using cage techniques within nursery areas, and at the scale of hectares using isolated populations of juvenile shore crabs in small nursery areas as mesocosms. The results were compared to habitat-specific distribution in the field.Shore crab megalopae and first instar juveniles (settlers) were distributed non-randomly among micro-habitats in the assessed nursery areas, with great densities in both mussel beds, eelgrass and filamentous algal patches (on average 114-232 settlers m⁻²), and significantly smaller densities on open sand habitats at all times (on average 4 settlers m⁻²). The same habitat-specific settlement pattern was found in cages where predators were excluded, suggesting that active habitat selection at settlement was responsible for the initial distribution. Older juveniles (second to ninth instar crabs) were also sparse on sand, but in contrast to settlers, were concentrated in mussel beds, which showed significantly greater densities than eelgrass and algal habitats. The cage experiment demonstrated a dynamic distribution of juvenile crabs. Young juveniles constantly migrated over open sand habitats (20 m or further) and colonized the experimental plots in a habitat-specific pattern that reflected the distribution in the field. This pattern was also found for very small crabs colonizing predator-exclusion cages, suggesting that selection of habitat by migrating juveniles caused the ontogenetic change in habitat use. Although post-settlement movements were great within nursery areas, juvenile dispersal at a regional scale appeared to be small, and the recruitment of juvenile shore crabs to the shallow bays occurred mainly through pelagic megalopae.Conservative estimates at the scale of whole nursery areas, based on migration trap data and field samples, indicated great mortality of settlers and early benthic stages of shore crabs. Results from the cage experiment suggest that predation by crabs and shrimp were responsible for the high settlement mortality. Both enclosed cannibalistic juvenile crabs and local predators on uncaged habitat plots caused significant losses of settlers in all habitats (on average 22% and 64% 3 day⁻¹, respectively). The effect of predators was highly variable between trials, but differed little between habitat types, and predation had no detectable proximate effect on juvenile distribution, despite the great losses. Small settlement densities on sand habitats in combination with a refuge at low prey numbers, and an aggregation of cannibalistic juvenile crabs in nursery habitats appear to decrease the effect of habitat-specific predation rates on the distribution of juvenile shore crabs. This study demonstrates that active habitat selection at settlement followed by a dynamic redistribution of young juveniles can be the proximate processes responsible for habitat-specific distribution of epibenthic juveniles, and indicate that predation represents a major evolutionary process reinforcing this behavior.     

Endogenous swimming rhythms of juvenile blue crabs, Callinectes sapidus, as related to horizontal transport

The blue crab Callinectes sapidus settles and metamorphoses in areas of aquatic vegetation in estuaries. Crabs in the first-fifth instar stages (J1-5) then emigrate from these areas by walking on the bottom or pelagic dispersal throughout estuaries. The present study was designed to characterize the timing of this migration pattern relative to the light-dark and tidal cycles. Field sampling in Pamlico Sound, NC, USA indicated that J4-5 juveniles were most abundant in the water column during the night. J4-5 juveniles were collected from Pamlico Sound in an area near Oregon Inlet that has semi-diurnal tides, a Mid-Sound area where tides are weak, and on the West side where regular tides do not occur. Crabs from all three sites had a circadian rhythm in which they swam up in the water column during the time of darkness in the field. Peak swimming consistently occurred at about 0300 h, but was not related to the timing of the tidal cycle. Similar results were obtained for juvenile crabs from an adjacent estuary having semi-diurnal tides. Dispersal at night reduces predation by visual predators, and allows early juvenile blue crabs to disperse planktonically from initial settlement sites.     

Invertebrate dispersal and habitat heterogeneity: Expression of biological traits in a seagrass landscape

Seagrass meadows harbour diverse faunal assemblages, but the relative importance of landscapes attributes, settlement processes and biological traits of individual species for recruitment patterns is poorly understood. To quantify the influence of habitat heterogeneity on larval, juvenile and adult post-larval dispersal, invertebrates (> 125 µm) were collected in benthic settlement traps at five occasions (June-August) in three habitats; continuous seagrass, seagrass patches and bare sand. The study was carried out by SCUBA diving in a subtidal (2.5 m depth) seagrass landscape dominated by Zostera marina L. in the Baltic Sea. Traps collected totally > 30 taxa, with non-significant effects of habitat on species richness and total abundance. Total number of invertebrates exhibited strong temporal variability, probably driven by wind-induced bedload and water column transport. Surprisingly, traps located in small (< 12 m²) patches contained on average almost twice as many individuals as traps located in the continuous vegetation. Dominating taxa such as nematodes, copepods, and oligochaetes were found in similar densities across the landscape. In contrast, location within the landscape had strong effects on bivalve settlement and redistribution patterns, resulting in significantly lower densities in continuous vegetation compared to patches and bare sand. A biological trait analysis indicated that semi-mobile taxa with annual protracted direct development, and short-distance dispersal are probably of higher importance for the community assembly process in this environment than long-distance larval dispersal. Results suggest that seagrass landscapes are highly dynamic environments, characterized by time and species-specific effects of landscape attributes on animal dispersal and recruitment. A conceptual model illustrating interactions between settling larvae and landscape heterogeneity is presented.     

Diet selectivity of juvenile blue crabs (Callinectes sapidus) in Chesapeake Bay

Shallow coves in Chesapeake Bay have abundant food and serve as nursery grounds for juvenile blue crabs. In this study, we examined the relationships between the diet of very small (4-40?mm CW) juvenile blue crabs and the benthic infauna in shallow, unvegetated nursery coves. We compared infauna in benthic samples with gut contents of juvenile blue crabs from six shallow coves in each of two sub-estuaries (Rappahannock and York Rivers) in Chesapeake Bay, Virginia, USA. Benthic communities differed depending on river and location, with abundant clams in upriver regions and abundant polychaetes in downriver regions. Juvenile crabs, like adults, appeared to be opportunistic feeders, with gut contents including clams, amphipods, polychaetes, small crustaceans, plant matter, and detritus. There was a positive relationship between polychaetes in the benthic samples and in crab guts, suggesting that juvenile crabs are opportunistic feeders on polychaetes in the benthos. Moreover, Ivlev's electivity index and foraging ratio showed that clams and polychaetes were selectively eaten at all locations. Alternatively, crabs selectively rejected amphipods. Crab densities corresponded positively with polychaete densities, which suggests that there may be bottom-up control of crab distributions and that food resources are important in nursery habitats.     

Impact Scenario for an Introduced Decapod on Arctic Epibenthic Communities

The intentional introduction of a species for the enhancement of stock or establishment of new fisheries, often has unforeseen effects. The red king crabs, Paralithodes camtschaticus, which was introduced into the Barents Sea by Russian scientists, has established a self-sustaining population that has expanded into Norwegian waters. As top benthic predators, the introduced red king crabs may have possible effects upon native epifaunal scallop (Chlamys islandica) communities. These benthic communities may be a source of prey species in late spring, when the red king crabs feed most intensively. Foraging rates (consumption, killing or severely damaging) of red king crab on native prey organisms were measured by factorial manipulation of crab density (0.5, 1.5 and 3 per m ²), size classes (immature, small mature, and large mature crabs), and by evaluating prey consumption after 48 h, in order to extrapolate a scenario of the likely impacts. Foraging rates of the red king crab on scallops ranged between 150 and 335 g per m² within 48 h. These rates did not change when crab density was altered, though an increased amount of crushed scallops left uneaten at the tank floor, were correlated with high density of small mature crabs. Foraging rate changed significantly with crab size. Consequently, the susceptibility of native, shallow water epibenthic communities to red king crab predation in the early life history stages, and during the post-mating/molting spring period, must be considered significant when foraging rates are contrasted with natural scallop biomass between 400 and 1200 g scallops per m².     

Assessing the potential benthic-pelagic coupling in episodic blue mussel (Mytilus edulis) settlement events within eelgrass (Zostera marina) communities

Coastal marine seagrass ecosystems are important nursery grounds for commercially and recreationally important species, and they serve as key settlement and recruitment sites for other species. We investigated several years (2001-2003) where episodic settlement events of blue mussels (Mytilus edulis) occurred in Barnegat Bay, NJ, USA. Population assessment indicated that blue mussels settled in eelgrass beds (Zostera marina) in late spring with peak densities exceeding 170,000 m⁻². Based on calculated filtration rates of M. edulis, we determined that for at least 53 days in 2001, the density and size distribution of M. edulis were sufficient to filter the water column volume in excess of twice a day, with maximum calculated filtration rates exceeding 8 m³ water m⁻² day⁻¹. While the settlement event in 2001 was very localized, in 2003, the settlement event was considerably more widespread throughout the bay, with maximum settling densities exceeding 175,000 individuals m⁻². Associated with these high densities, maximum calculated filtration rates exceeded 15 m³ water m⁻² day⁻¹. This filtration potential may have impeded the localized development of a brown-tide (Aureococcus anophagefferens) bloom in 2001, which occurred in other regions of the bay, but the widespread settlement event seen in 2003 may have impeded the development of any brown-tide blooms in Barnegat Bay during that summer. The decline in mussel densities throughout the summer may be a result of elevated water temperatures in this back bay, but at one site, the high settlement of M. edulis was followed by a substantial migration (>40 individuals m⁻²) of small sea stars (Asterias forbesii). In 2001, A. forbesii was a significant factor in reducing M. edulis density by the end of the summer at the Barnegat Inlet site and a community level assessment showed significant positive correlations between mussel aggregations and sea star densities (r=0.68-0.73, P<0.001). At this same site in 2003, the sea stars were again present in high densities (26 m⁻²) and were a potential mechanism for mussel decline. In other regions of the bay, sea star densities are very low, but numerous other predatory species exist, including blue crabs (Callinectes sapidus), green crabs (Carcinus maenus), spider crabs (Libinia spp.), and several Xanthid crabs. Given the high mussel densities seen in this study and the considerable predation by sea stars and other benthic predators, the benthic-pelagic coupling which these mussels provide in this system contributes to the high secondary production in these grass beds.     

Interactive effects of episodic hypoxia and cannibalism on juvenile blue crab mortality

We hypothesized that as the spatial extent of hypoxic bottom water increased, (1) adult blue crab predator densities would increase in shallow habitats as they avoided hypoxia, and that (2) juvenile blue crabs, which use shallow unvegetated habitat as a predation refuge from adult conspecifics, would experience increased mortality rates during crowding by cannibalistic adult blue crabs. These hypotheses were tested along a depth gradient of sandy-mud shoreline in the Neuse River Estuary (NRE), North Carolina, USA using a combination of (1) hydrographic measurements to characterize the spatial extent of hypoxia, (2) beach seines to quantify the density of adult blue crab predators in relatively shallow water as a function of 1, and (3) tethering experiments to quantify relative rates of predation on juvenile blue crabs as a function of 1 and 2. During our seven tethering experiments, the NRE study site experienced a range of DO scenarios including normoxia, chronic hypoxia, and hypoxic upwelling. No known predators of juvenile blue crabs, other than adult conspecifics, were collected in any of our shallow-water seines. During the transition from normoxia to chronic hypoxia, blue crab predator densities in shallow refuge habitats increased 4-fold, and relative mortality rates of juvenile blue crabs in shallow habitats increased exponentially with the density of adult conspecifics. Conversely, during hypoxic upwelling events, the density of adult blue crabs in shallow water declined, which may explain why the relative mortality of juvenile crabs did not increase significantly with the increasing spatial extent of hypoxia. Thus, juvenile blue crabs may be relatively safe from adult conspecifics during hypoxic upwelling events, but not during chronic hypoxia. These experimental results highlight the need to consider the effects of dynamic water quality on mobile consumers emigrating from degraded habitats when considering indirect trophic impacts beyond the immediate area of impact.     

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

Processes regulating early post-settlement habitat use in a subtidal assemblage of brachyuran decapods

In highly mobile animals post-settlement dispersion of juveniles can strongly influence the observed patterns of abundance and distribution. To explore the relative importance of factors regulating the use of habitat by crabs we performed a multi-species manipulative experiment in a subtidal environment of the central Chilean coast. First, demographic patterns were established by performing a year-round crab survey in three discrete and well-known subtidal crab habitats: (1) algal turf, (2) cobbles and (3) shell hash. Second, habitat preferences were experimentally evaluated using concrete trays that were filled with different substrate types that simulate natural habitats. Settlement and recruitment rates were estimated from experimental trays that were left in the field and surveyed after 2 weeks (complete experiment was repeated 7 times throughout 1 year). Third, mortality, due to predation, was assessed by covering 50% of the trays with a 4-mm mesh-size screen that excluded large predators (i.e., fishes, shrimps). Fourth, habitat colonization rates were evaluated by quantifying the arrival, into open trays, of large juveniles (secondary dispersal). The most abundant species in this system (Paraxhantus barbiger, Cancer setosus, Taliepus dentatus and Pilumnoides perlatus) displayed clear habitat preferences at the time of settlement, evidenced by differences in density of recruits among habitats. Recruitment regulation by predation seemed to explain the observed patterns in only one case. For most species, however, evidence of ontogenetic change in the use of habitat, through active habitat redistribution by large juveniles, was detected. Thus, secondary dispersal among habitats seems to outweigh the influence of megalopae's habitat selection and post-settlement mortality as responsible for the observed demographic patterns.     

Spatial distribution of the upside-down jellyfish Cassiopea sp. within fringing coral reef environments of the Northern Red Sea: implications for its life cycle

The zooxanthellate mangrove jellyfish Cassiopea sp. represents a prominent invasive species and a potential bioindicator for nutrient monitoring in coral reefs. However, information about its spatial distribution in combination with abundance, habitat specificity and life cycle elements is barely available. This study, therefore, presents the results of field surveys conducted within four different benthic habitat types (coral reef, seagrass meadow, reef-sand transition and sand flat) in the Northern Red Sea. Cassiopea sp. exhibited a highly patchy distribution within the entire study area with mean abundance of 1.6 ± 0.3 animals m⁻² and benthic coverage of 3.2%. Within coral reef habitats, maximum abundance of up to 31 animals m⁻² and benthic coverage of up to 20% were detected. Additionally, this study revealed that 65% of all observed Cassiopea specimens were associated with the commensalistic crustacean mysid Idiomysis tsurnamali. Cassiopea abundance and size as well as association patterns with mysids differed between most of the surveyed habitats. In summary, the findings of the present study (1) characterize Cassiopea as one of the key organisms in investigated benthic habitats, (2) indicate active habitat selection by the jellyfish and (3) may hint to an unexplored life cycle of Cassiopea with central role of seagrass meadows providing cues for larval settlement and metamorphosis in the absence of mangroves.     

The synergism between hydrocarbon pollutants and UV radiation: a potential link between coastal pollution and larval mortality

Coastal, benthic invertebrates with complex life history strategies are exposed to stage- and habitat-specific selective forces. In the coastal environment, benthic adults are exposed to polycyclic aromatic hydrocarbon pollutants (PAHs) due to their proximity to human activities (shipping, urbanization, and industrialization). Benthic invertebrates produce lipid-rich eggs or larvae that absorb PAHs from polluted estuaries and coastal waters. The larvae of many coastal invertebrates move offshore following release from benthic adults. During development in offshore waters, larvae of some species are exposed to relatively high levels of ultraviolet (UV) radiation. Marine organisms vary in their tolerance to PAHs and UV radiation. The purpose of this study was to examine the effects of the sequential exposure of the larvae of marine crabs to PAHs and UV radiation.Using laboratory experiments, the larvae of four crab species were exposed to PAHs and UV radiation. There was a significant synergistic effect of exposure to PAH (fluoranthene or pyrene) and UV radiation on larvae of the spider crab (Libinia dubia), the stone crab (Menippe adina) and the mud crab (Panopeus herbstii). Larvae of blue crabs (Callinectes sapidus) were exposed to PAHs and UV radiation in both laboratory and solar UV experiments. Significantly higher mortality occurred for C. sapidus larvae using either type of UV-artificial or solar.Larvae of coastal invertebrates with complex life history strategies are susceptible to the combined effects of PAHs and UV radiation. In this study, the exposure of crab larvae to PAHs and UV radiation resulted in mortality to crab larvae using laboratory and solar UV experiments. There were no effects on larval crab mortality due to PAH or UV radiation independently but mortality was as high as 100% when both factors were present.     

Facultative mutualism between an herbivorous crab and a coralline alga: advantages of eating noxious seaweeds

Because encrusting coralline algae rely on herbivory or low light levels to prevent being overgrown by competitively superior fleshy algae, corallines are relatively rare in shallow areas with low rates of herbivory. In contrast to this general trend, the branching coralline alga Neogoniolithon strictum occurs primarily in shallow seagrass beds and along the margins of shallow reef flats where herbivory on macrophytes is low. This alga apparently persists in these habitats by providing refuge to the herbivorous crab Mithrax sculptus at mean densities of 1 crab per 75 g of algal wet mass. When crabs were removed from some host corallines, hosts without crabs supported 9 times the epiphytic growth of hosts with crabs after only 30 days. Crabs without access to a coralline alga were rapidly consumed by reef fishes, while most of those tethered near a host alga survived. These results suggest that the crabs clean their algal host of fouling seaweeds and associate with the host to minimize predation. However, to effectively clean the host, the crab must consume the wide array of macroalgae that commonly co-occur with coralline algae in these habitats, including chemically defended species in the genera Halimeda, Dictyota, and Laurencia. Crabs did readily consume these seaweeds, which were avoided by, and are chemically defended from, herbivorous fishes. Even though crabs readily consumed both Halimeda and Dictyota in whole-plant feeding assays, chemical extracts from these species significantly reduced crab feeding, suggesting that factors other than secondary chemistry (e.g., food value, protein, energy content), may determine whole-plant palatability. Having the ability to use a wide variety of foods, and choosing the most profitable rather than the least defended foods, would diminish foraging time, increase site fidelity, and allow the crab to function mutualistically with the host alga. Despite the obvious benefit of associating with N. strictum, M. sculptus did not prefer it over other habitats offering a structurally similar refuge, suggesting that these crabs are not N. strictum specialists, but rather occupy multiple habitats that provide protection from predators. Structurally complex organisms like N. strictum may commonly suppress competitors by harboring protective symbionts like M. sculptus. It is possible that diffuse coevolution has occurred between these two groups; however, this seems unlikely because both herbivore and host appear to respond most strongly to selective pressures from predators and competitors outside this association.     

Post-settlement effects of habitat type and predator size on cannibalism of glaucothoe and juveniles of red king crab Paralithodes camtschaticus

Postlarval (glaucothoe) and juvenile (first crab stage, C1) red king crab Paralithodes camtschaticus actively select structurally complex substrata for settlement. Such habitats may provide them with shelter from predation during critical early stages. We tested this hypothesis by placing glaucothoe and juvenile crab in aquaria with or without natural or artificial habitats, and with or without predators (1-3-year-old red king crab) of two different sizes. Predators caused increased mortality of glaucothoe, but predator size, habitat presence and habitat type had no effect on survival. Predators caused significant mortality of C1 crabs in the absence of habitat, and mortality was inversely related to predator size. Density of glaucothoe on habitats was similar with or without predators, but density of C1 crab on habitats was higher than that of glaucothoe, and increased in the presence of large predators. Active selection for complex substrata by settling glaucothoe does not reduce cannibalism, but may pre-position them for improved survival after metamorphosis. In contrast, juvenile crabs modify their behavior to achieve higher densities in refuge habitats, which tends to dampen the effect of predation. These survival strategies may have evolved to compensate for the greater risk of predation in open habitats.     

Trophic models of four benthic communities in Tongoy Bay (Chile): comparative analysis and preliminary assessment of management strategies

Steady-state trophic flow models of four benthic communities (seagrass, sand-gravel, sand and mud habitats) were constructed for a subtidal area in Tongoy Bay (Chile). Information of biomass, catches, food spectrum and dynamics of the commercial and non-commercial populations was used and the ECOPATH II software of Christensen and Pauly [Ecol. Modell. 61 (1992a) 169] was applied. The sea star Meyenaster gelatinosus and the crabs Cancer polyodon, C. porteri and Paraxanthus barbiger were found to be the most prominent predators in the benthic system. The scallop Argopecten purpuratus as well as other bivalves represented the principal secondary producers in the seagrass, sand-gravel and sand habitats, while the Infauna dominated the mud habitat. The highest total biomass and system throughput (33579.3 t/km²/year) was calculated for the sand-gravel habitat. The sand habitat had a negative net system production due to the amount of primary production imported from deeper waters to satisfy the food requirements of the large beach clam (Mulinia sp.) populations. The mean trophic level of the fishery varied between 2.06 (sand-gravel) and 3.92 (sand) reflecting the fact that the fishery concentrates on primary producers (i.e. algae and filter feeding), and on top predators (i.e. snails and crabs). Fishery is strongest in sand-gravel habitat, where annual catches amount to 122.05 g/m². Low values of the relative Ascendency (A/C) (from 27.4 to 32.7%) suggest that the systems analysed are immature and highly resistant to external perturbations. Manipulations of the input data for the exploited species suggest that seagrass and sand-gravel habitats have a potential for a ∼3 times higher than the present production of scallops and the red algae Chondrocanthus chamissoi. Preliminary results of Mixed Trophic Impacts (MTI) analysis suggest that any management policy aimed at a man-made increase in the standing stocks of A. purpuratus and Ch. chamissoi in seagrass and sand-gravel habitats, and a removal of the seastar M. gelatinosus in the seagrass habitat appears justified.     

Small scale variability of benthic assemblages: biogenic neighborhood effects

In this study, patterns of community development were investigated within vs. outside 'habitats'. These habitats represented five different monospecific assemblages of one of the following species: the brown alga Fucus serratus, the red alga Delesseria sanguinea, the green alga Enteromorpha intestinalis, the seagrass Zostera marina and the blue mussel Mytilus edulis. Natural assemblages were allowed to develop on paired artificial substrata-separated by ca. 1 m-within (treatment) vs. outside (control) of habitats. The same colonizer species settled on treatment and control substrata for given habitats. However, after 5 months of settlement and post-settlement dynamics, their proportional abundance and the structure of treatment and control assemblages differed in many instances. Variability among replicates of a given treatment, seperated by up to 50 m, was large, indicating a patchy spatial distribution of organisms. Despite this spatial heterogeneity among within-treatment replicates, analysis of similarity revealed that in most instances significantly different assemblages developed between treatments on a small spatial scale depending on whether substrata were positioned within as compared to outside a given habitat.Consequently, the algae, seagrass or mussels constituting a habitat seem to control the structure of the benthic assemblage developing in their vicinity by one or more possible mechanisms: reduction of larval advection, exudation of metabolites that influence settlement and/or post-settlement survival, and/or-in the case of mussel assemblages-predation on larvae.In addition to spatial variability in larval supply, stochasticity in succession, substratum heterogeneity, competition and predation effects, this investigation reveals the potential of a further assemblage structuring factor: the impact of neighboring organisms.     

The intertidal mudflats of Barr Al Hikman,Sultanate of Oman,as feeding,reproduction and nursery grounds for brachyuran crabs

Brachyuran crabs are an important ecological and economical, yet often unstudied aspect of intertidal mudflats of the Arabian Peninsula. Here we provide baseline density estimates of crabs at the relatively pristine intertidal mudflats of Barr Al Hikman (Sultanate of Oman) and provide information on their life cycle and habitat preference. Across the winters of 2012–2015 crabs were sampled on a grid covering the entire intertidal depth gradient. 29 species were found and average densities varied between 12 and 54 crabs/m². Deposit-feeding and herbivorous crabs were the most abundant species across all winters. Size frequency data and the presence of ovigerous females show that most crabs species reproduce in the intertidal area. P. segnis, the most important crab for local fisheries, was found to use the intertidal area as a nursery ground. We analysed the relationships between the two most abundant crab species, Macrophthalmus sulcatus and Thalamita poissonii and the environmental variables: seagrass density, tidal elevation, median grain size and sediment depth using Random Forest models. The predictive capacity of the models and the relative importance of the environmental predictors varied between years, but crab densities in general were positively associated with seagrass density, presumably because seagrass offers feeding habitat.

     

Mechanism for the small-scale movement of carbon among estuarine habitats: organic matter transfer not crab movement

In theory, carbon is highly mobile in aquatic systems. Recent evidence from carbon stable isotopes of crabs (Parasesarma erythrodactyla and Australoplax tridentata), however, shows that in subtropical Australian waters, measurable carbon movement between adjacent mangrove and saltmarsh habitats is limited to no more than a few metres. We tested whether the pattern in crab δ¹³C values across mangrove and saltmarsh habitats was explained by crab movement, or the movement of particulate organic matter. We estimated crab movement in a mark–recapture program using an array of pitfall traps on 13 transects (a total of 65 traps) covering an area of 600 m² across the interface of these two habitats. Over a 19-day period, the majority of crabs (91% for P. erythrodactyla, 93% for A. tridentata) moved <2 m from the place of initial capture. Crab movement cannot, therefore, explain the patterns in δ¹³C values of crabs. δ¹³C values of detritus collected at 2-m intervals across the same habitat interface fitted a sigmoidal curve of a similar form to that fitting the δ¹³C values of crabs. δ¹³C values of detritus were 2–4‰ more depleted in saltmarsh (−18.5±0.6‰), and 4–7‰ more depleted in mangroves (−25.9±0.1‰) than δ¹³C values of crabs recorded previously in each habitat. Assimilation by crabs of very small detrital fragments or microphytobenthos, more enriched in ¹³C, may explain the disparity in δ¹³C values. Nevertheless, the pattern in δ¹³C values of detritus suggests that crabs obtain their carbon from up to several metres away, but without themselves foraging more then a metre or so from their burrow. Such detailed measurements of carbon movement in estuaries provide a spatially explicit understanding of the functioning of food webs in saltmarsh and mangrove habitats.