Temporal and ontogenetic shifts in habitat use of juvenile Pacific cod (Gadus macrocephalus)

Habitat use of age-0 and age-1 juvenile Pacific cod (Gadus macrocephalus) was examined in coastal regions in Kodiak Alaska over daily, seasonal and annual scales. Catch data indicated highly variable recruitment to nursery areas, but a strong separation of distribution by depth among age groups. Age-0 cod were most abundant in the shallows (<3 m) whereas age-1 cod were typically found in depths (9.0-13.5 m). In comparison, age-1 saffron cod (Eleginus gracilis), another highly abundant gadid in the region, were found in shallower depths where age-0 cod often resided. Age-1 cod Pacific cod made diel lateral movements, moving into shallow regions at night where they co-occurred with age-0 cod to a greater extent. Laboratory light-gradient experiments indicated that age-0 cod tolerated intense lighting (~ 20-80 µE m^− 2 s^− 1) typical of shallow water regions whereas larger age-1 Pacific cod strongly avoid bright light given the choice. However, while diet data indicate age-1 cod of both species are moderately piscivorous (3% saffron cod; 16% Pacific cod), we found no direct evidence of predation on smaller conspecific cod, possibly due to the low densities of age-0 cod in the year of the diet study. Together, these data suggest that coastal regions continue to serve a nursery function beyond the 1st year of development for juvenile Pacific gadids, and that small-scale temporal and depth partitioning in these regions is a mechanism by which varying cod species and age classes co-occur.     

Conformation and stability of elastase from Atlantic cod, Gadus morhua

Metal binding and conformational stability characteristics of psychrophilic elastase (ACE) from Atlantic cod (Gadus morhua) has been investigated. Chelation to Ca²⁺ was found to be important for maintaining the biologically active conformation and for the thermal stability of the enzyme. However, presence of metal ions such as Zn²⁺, Fe³⁺ and Cu²⁺ was found to inhibit its hydrolytic activity and so did the chelating agent EDTA. Both pH and guanidinium chloride induced denaturation of the enzyme was followed by monitoring the changes in the tryptophan fluorescence. ACE exhibited a simple two-state unfolding pattern in both acidic and basic conditions with the midpoint of transition at pH values 4.08 and 10.29, respectively. Guanidinium chloride and heat induced denaturation of the enzyme was investigated at two pH values, 5.50 and 8.00, wherein the enzyme possesses similar tertiary structure but differ in its hydrolytic activity. Guanidinium chloride induced denaturation indicated that the enzyme unfolds with a C_m of 1.53 M at pH 8.0 and a ΔG_H2O of 6.91 kJ mol⁻¹ (28.65 J mol⁻¹ residue⁻¹) which is the lowest reported for psychrophilic enzymes investigated till-date. However, at pH 5.50, ΔG_H2O value is slightly lowered by 0.65 kJ mol⁻¹ consistent with the observed increase in the apparent quenching constant obtained with acrylamide. On the other hand, increase in T_m by 38.45 °C was observed for the enzyme at acid pH (5.50) in comparison to the heat induced unfolding at pH 8.0. The increase in the apparent T_m has been attributed to the possible weak intermolecular association of the enzyme molecules at moderately high temperatures that is favoured by the increase in the accessible surface area / dynamics under acidic conditions. The stability characteristics of ACE have been compared with the available data for mesophilic porcine pancreatic elastase and possible mechanism for the low temperature adaptation of ACE has been proposed.     

Seasonal variations in seminal plasma and sperm characteristics of wild-caught and cultivated Atlantic cod, Gadus morhua

The objective was to investigate changes, throughout the spawning season, in body size attributes and quantitative semen characteristics of wild-caught and cultivated Atlantic cod, Gadus morhua L. Sperm velocity increased significantly throughout the spawning season of cod from both origins. Curvilinear velocity (VCL; 30 sec post-activation) increased from 78.9 ± 6.5 to 128.2 ± 6.5 μm/sec (mean ± SEM) between the beginning and end of the spawning season, respectively, for wild-caught cod, whereas for cultivated fish, it increased from 26.6 ± 2.4 to 48.9 ± 3.1 μm/sec between January and March. Spermatocrit did not undergo a significant seasonal change in wild-caught cod but did thicken for cultivated cod (24.6 ± 4.2% in January to 40.5 ± 4.4% in April; P < 0.01). Sperm head area, perimeter, length, and width declined significantly at the end of the spawning season of cod from both origins (all P values < 0.01). Seminal plasma osmolality and Na⁺ ion concentration followed a dome-shaped function through the spawning season for both wild-caught and cultivated cod (P < 0.05). For cultivated cod, seminal plasma pH was significantly lower at the start of the spawning season (P < 0.001), whereas Ca²⁺ increased then decreased (P < 0.05). Body size attributes, spermatocrit, and seminal plasma constituents had significant relationships with sperm activity variables. These relationships varied as a function of time post-activation, month, and fish origin. Our findings may be used to (i) assess spermiation stage without killing males; (ii) optimize semen collection for hatchery production; (iii) characterize the potential impact of farming on sperm quality; and (iv) improve success of sperm cryopreservation and short-term storage.     

Eelgrass patch size and proximity to the patch edge affect predation risk of recently settled age 0 cod (Gadus)

Postsettled age 0 Atlantic cod (Gadus morhua) seek refuge from predation in eelgrass (Zostera marina) habitat within shallow, coastal nurseries. Laboratory and field experiments have demonstrated that predation risk on small fish is reduced in habitats of greater structure compared to less complex or barren environs. To determine if predation risk is linked to the areal extent of eelgrass coverage, we tested the hypothesis that predation risk of age 0 cod decreases with increasing eelgrass patch size. Predation on tethered age 0 cod (G. morhua and G. ogac) was quantified in eelgrass patches (1-80 m²) at three sites in northeastern Newfoundland, Canada, during September and October 1999 and 2000. Based on evidence of edge effects in terrestrial landscapes, we also tested the hypothesis that predation is elevated at ecotones separating eelgrass from adjacent barren mud substrate. We examined predation at the edge (i.e., 0 m) and both 5 and 10 m from this edge into and away from eelgrass cover along an 18 m long barren mud-gravel and eelgrass boundary, at two sites. Logistic regression analysis showed that the risk of predation, as measured by the odds ratio, increased with area over a small range of patch sizes (1-35 m² in 1999). When the study was extended to a wider range of patch sizes (1-80 m²) in 2000, a parabolic relationship emerged, with patches on the order of 25 m² providing the least safety and the largest patches (80 m²) providing the most safety. Predation on tethered cod was highest at the edge of eelgrass patches, compared with barren and eelgrass locations; predation generally decreased with distance from the eelgrass boundary. Our results are consistent with the hypothesis that predators are drawn to large patches of eelgrass because of increased prey numbers, that predators increase their success by searching edges, and that this results in greatest predation risk to prey in isolated patches of intermediate size.     

Identification of lipids in the cuticle of the parasitic nematode Anisakis simplex and the somatic tissues of the Atlantic cod Gadus morhua

The main aim of this work was to assign the cuticular lipids identified in a parasitic nematode and to distinguish those originating from its host. The hypothesis that long-chained fatty acids and sterols are imported by the parasite in the absence of certain enzymes was also tested. The organisms (Anisakis simplex and Gadus morhua) were extracted in petroleum ether and dichloromethane. Matrix-assisted laser desorption time-of-flight mass spectrometry (MALDI-TOF) was used to identify unknown components, and electrospray ionization mass spectrometry (ESI/MS) to verify recognized groups of lipids. The lipid classes identified in the surface layer were free saturated and unsaturated fatty acids, triacylglycerols, sterols and non-polar sphingolipids (ceramides, sphingoid bases). The most abundant fraction consisted of fatty acids. The predominant saturated acids were tetradecanoic acid in the petroleum ether extract of A. simplex, hexadecanoic acid in the dichloromethane extract of A. simplex, and also the polyunsaturated octadecahexaenoic and octadecatrienoic acids in both extracts of the parasitic nematode. The mass spectrum revealed the presence of fatty acids with different numbers of carbons, and with odd and even numbers of unsaturated bonds. The MALDI-TOF mass spectrum also identified triacylglycerols (TAGs). The dominant short-chain TAGs were CoCoCy:₁, CoCoPg and Bu0:0B:₆. The majority of TAGs were found in the ether and dichloromethane extracts of A. simplex. Sterols were the least common class of lipids found in the nematode extracts; most likely, this is the fraction that is entirely incorporated from the host organism because of the parasite’s inability to synthesize them. MALDI-TOF also identified non-polar sphingolipids - ceramides and sphingoid bases. The signals due to N-octanoyl-d-erythro-octasphinganine (m/z 288.3) and N-tetranoyl-d-erythro-tetradecasphinganine (m/z 316.4) were dominant on the mass spectra; quite a large number of short-chain non-polar sphingolipids were also identified.     

Effect of somatic and otolith growth rate on stable isotopic composition of early juvenile cod (Gadus morhua L) otoliths

The relative amounts of the stable isotopes of carbon and oxygen in fish otoliths can be used to reveal the environmental history experienced by the fish. This requires that the relative amounts of the isotopes are deposited in equilibrium with the surrounding environment, or that the offset from this equilibrium is known and can be quantified. It is known that carbon isotopes in biogenic carbonates are a mixture of carbon from the seawater and metabolically derived carbon, but the effect of the somatic growth rate of the fish is still unclear. The possible effect of otolith growth rate and fractionation of both carbon and oxygen isotopes are also not established. We carried out a controlled laboratory experiment where we reared cod (Gadus morhua L.) larvae and early juveniles at two temperatures (6 and 10 °C) and generated different growth rates within each temperature by manipulation of prey levels. The otoliths of the resulting fish were analysed for carbon and oxygen isotopes. We found no effect of otolith precipitation rate on fractionation of either carbon or oxygen isotopes. However, there was a depletion of ¹³C in the otoliths of fish with elevated metabolism. The proportion of metabolically derived carbon in the otoliths was estimated to be 28-32%. Our results suggest that measurements of oxygen isotopes in otoliths can be a reliable tool to estimate ambient temperature since the oxygen isotopes seem to be deposited in the otoliths independently of kinetic and metabolic effects. Fractionation of carbon isotopes in otoliths on the other hand can give valuable insight into metabolism and feeding pattern of fish.     

The effect of growth rate,size, and season on oocyte development and maturity of Atlantic cod (Gadus morhua L.)

The effect of growth rate, body weight, age, and season on ovarian development and maturation was investigated for Atlantic cod (Gadus morhua L.) reared in the laboratory over 10 months, for each of two consecutive years, 1978–1980. Cod were also collected from the Gulf of St. Lawrence, the Scotian Shelf, Georges Bank, the Flemish Cap, and the N.E. Newfoundland Shelf.The state of maturity was recognized by oocyte size. Stage I oocytes did not vary in size with growth rate, season, age, or maturity, while the size of stage II oocytes was positively correlated with maturity and negatively correlated with maximum stage of development achieved. Ovarian wall thickness was positively correlated with age and maturity.The frequency distribution of stage I and II oocytes distinguished the state of maturation, with cod that would mature by the next spawning season having a minimum of 20% ($\text{x}\text{?}\text{= 42\%}$) of their oocytes at stage II or greater level of development.Maturing 3-yr-old cod had greater life specific growth rates than immature 3-yr-olds, but growth rates during the third year itself were not significantly different. A hypothesis of a three-part density-dependent mechanism controlling fecundity is postulated. Future reductions in partial recruitment and total fecundity are predicted for the Gulf of St. Lawrence cod stock based on calculated growth rates for Gulf cod in 1979.     

Cryopreservation of Atlantic cod Gadus morhua L. spermatozoa: Effects of extender composition and freezing rate on sperm motility, velocity and morphology

Broodstock selection programs are currently underway for Atlantic cod (Gadus morhua). To complement and further these selection programs we need to develop sperm cryopreservation procedures. This will allow genomic DNA from males from selected individuals or stocks to be frozen and conserved in perpetuity. In our study we used a full factorial ANOVA design to examine the effects of diluent (Mounib’s sucrose-based diluent, Hanks’ Balanced Salt Solution, Mounib’s sucrose-based diluent + hen’s egg yolk, and Hanks’ Balanced Salt Solution + hen’s egg yolk), cryoprotectant (propylene glycol, dimethyl sulphoxide, and glycerol), and freezing rate (−2.5, −5.0, −7.5, and −10.0 °C/min) on motility of cod frozen-thawed sperm. Sperm velocity and morphometric analyses of sperm heads and flagella were also assessed. We found that sperm motility-recovery index was strongly influenced by the presence of higher-order interactions of the factors we tested. The best cryoprotection used diluents that contained hen’s egg yolk. Generally, extenders containing propylene glycol yielded higher post-thaw sperm motilities than those with dimethyl sulphoxide or glycerol. In comparison to sperm from other frozen-thawed extenders, sperm from extenders supplemented with propylene glycol had significantly higher curvilinear velocity. Cryopreservation showed no impact on sperm head morphology parameters, however, considerable damage to frozen-thawed sperm flagella was observed. We believe that our experimental/statistical approach and our results add significantly new information to the study of semen biology/cryobiology in fishes. Our findings are also highly relevant to the development of cod mariculture and for aiding in conservation efforts of this very important marine species.     

Ontogenetic changes in the critical swimming speed of Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin) larvae and the role of temperature

Most studies on behavioural contributions to dispersal and recruitment during early life history stages of fishes have focused on coral reef species. For cold ocean environments, high variation in seasonal temperature and development times suggest that parallel studies on active behaviour are needed for cold-water species. Thus, we examined the critical swimming speed (U_crit) of marine fish larvae from 2 contrasting species: Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin), a pelagic and bottom spawner respectively. Within-species comparisons showed that sculpin reared at 6 °C had lower initial U_crit values, but a faster U_crit increase through development compared with 3 °C conspecifics, ultimately resulting in faster critical swimming speeds at metamorphosis (10.5 vs. 9.1 cm·s^− 1). In contrast, although cod larvae reared at 10 °C were faster swimmers at first feeding than 6 °C fish, temperature differences were absent after the first week. These results show that temperature influences the trajectory of larval critical swimming speed development, but that the relationship is species-specific. Although 6 °C sculpin and cod of similar length had equivalent U_crit values, the smaller size of cod at hatch (5.3 vs. 10.8 mm for sculpin) resulted in much lower age-specific U_crit values for cod. These data have significant implications for how swimming activity of the two species might affect dispersal, particularly in the first few weeks post-hatch. Overall, our data suggest that temperature during larval development influences the swimming capacity of cold-water marine fishes, and has important ramifications for biophysical models of dispersal.     

Hypoxic avoidance behaviour in cod (Gadus morhua L.): The effect of temperature and haemoglobin genotype

Hypoxia can influence fish growth, survival and on larger scales, population structure. These effects may be influenced by water temperature, and may vary intra-specifically with genotype. In Atlantic cod (Gadus morhua L.), the two haemoglobin homozygotes (Hb-I?11 and Hb-I?22) vary in oxygen affinity at different temperatures, which is thought to correspond to variation in hypoxia tolerance. We therefore tested if hypoxic avoidance behaviour in cod 1) depends on ambient temperature and 2) is modified by haemoglobin genotype. In a laminar flow choice box, we subjected juvenile cod to an initial phase of non-escapable hypoxia, and a subsequent recovery phase, where one habitat was kept at 20% O₂ saturation while the other was raised in steps to full saturation. The experiment was performed at 5 and 15 °C with Hb-I?11 and Hb-I?22 cod. Cod responded to inescapable hypoxia by reducing their overall swimming speed and then, at the initial levels of the recovery phase, avoiding the most hypoxic habitat, irrespective of temperature or genotype. Fish recovered quickly as O₂ levels increased, as evidenced by increased swimming speed and time spent in the most hypoxic habitat. The avoidance response depended strongly on temperature: the relative reduction in speed and avoidance of the most hypoxic habitat was more pronounced at 15 than at 5 °C. During the recovery phase, stressed fish initially maintained a higher swimming speed in the most hypoxic habitat. However, as O₂ increased, swimming speed in both habitats converged. This point of convergence occurred at a lower O₂ saturation at 5 °C. Fish ventilation rate in inescapable hypoxia was also higher at 15 °C. Haemoglobin genotype did not influence either ventilation rates or the nature of the hypoxic avoidance response at either temperature, but Hb-I?11 cod swam faster than Hb-I?22 cod in normoxia at 15 °C. Our results indicate that increased temperature limits the ability of cod of both haemoglobin genotypes to exploit hypoxic habitats. This may have negative future consequences for coastal cod stocks in light of increasing global temperatures and eutrophication in coastal waters.     

Attachment of the juvenile queen scallop (Aequipecten opercularis (L.)) to maerl in mesocosm conditions; juvenile habitat selection

Substratum un-acclimated juvenile queen scallops (Aequipecten opercularis) (<18 mm and 18-30 mm shell height) were released in equal numbers onto pristine live maerl (PLM), impacted dead maerl (IDM), gravel and sand in choice chambers. Their habitat selection was monitored over a 4-day period in control and predator treatments (utilising Asterias rubens L. and Carcinus maenas (L.)). Microhabitat use of PLM by juvenile queen scallops and the presence of cues in live maerl were also investigated.In control and predator treatments juvenile queen scallops were observed to attach preferentially to PLM than IDM, gravel or sand. Juvenile queen scallops were observed to maintain a more exposed attachment site in the absence of predators but sought refuge within and between maerl nodules in the presence of both predators. Smaller queen scallops (<18 mm shell height) were more efficient at utilising maerl thalli as a refuge. Juvenile A. opercularis showed hierarchical cue responses mediated by predator presence, i.e. responding favourably to a factor associated with live maerl presence irrespective of heterogeneity in the absence of predators but favourably to higher maerl heterogeneity in their presence. If they also preferentially attach to PLM in the field, at some sites where PLM grounds cover large areas, they may thus be considered to constitute ‘nursery areas’. Habitat attachment preference appears to be predetermined and not a result of localised predator avoidance; however, habitat usage changes in the presence of predators. Maerl beds have been shown to be easily damaged by scallop dredging in Scotland and if such nursery areas are being destroyed extensively in the field, this could damage recruitment to localised adult populations.     

Responses of the black sea urchin Tetrapygus niger to its sea-star predators Heliaster helianthus and Meyenaster gelatinosus under field conditions

We ran field experiments to examine the responses of the black sea urchin Tetrapygus niger to predatory sea stars. Trials involving simulated attacks (one or several arms of a sea star being placed on top of half the urchin) showed that the urchin differentiated between the predatory sea stars, Heliaster helianthus and Meyenaster gelatinosus, and a non-predatory sea star, Stichaster striatus, and showed almost no response to a sea star mimic. We further compared the responses of the urchin to different threat levels presented by the two predatory sea stars. The highest threat level was a simulated attack, then mere contact, and subsequently sea stars being placed at different distances from the urchin. All urchins responded to simulated attacks and contact with both sea stars. The proportion responding decreased with distance and more rapidly in trials with H. helianthus (0% at a distance of 30 cm) than with M. gelatinosus (33% at a distance of 50 cm). At each of the threat levels where there was a response to both sea stars, the urchins responded more rapidly to M. gelatinosus than to H. helianthus. In a third experiment where a predatory sea star was added to a circular area (1-m diameter) in which either 4-8 or 11-19 undisturbed urchins were present, the urchins fled the area more rapidly when the added sea star was M. gelatinosus, but the rate of fleeing did not vary with density, as might occur if there was communication among urchins using alarm signals. Our observations suggest that M. gelatinosus presents a stronger predatory threat than H. helianthus. This corresponds to field observations showing that the urchins are more frequently consumed by M. gelatinosus. These are the first field experiments demonstrating distance chemodetection by a marine invertebrate under back-and-forth water flow from wave activity.     

Small- and large-scale effect of the SW Atlantic burrowing crab Chasmagnathus granulatus on habitat use by migratory shorebirds

Here we address the question of whether the presence of the burrowing crabs Chasmagnathus granulatus affects small- and large-scale habitat use by migrant shorebirds. This crab is the dominant species in soft bare sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 42°S). They generate very extensive burrow beds in soft bottom intertidal areas. Our information shows that this burrowing crab affects the small-scale habitat use by shorebirds, given that shorebirds never walk through the funnel-shaped entrances of burrows. Given that crab burrow entrances occupy up to 40% of the intertidal area, there is a large decrease of available shorebird habitat in crab beds, restricting their activity to the spaces between the burrows. The southern migratory shorebird Charadrius falklandicus maximize the use of these areas by foraging closer to the burrows than the other bird species. Neotropical migrants, such as Calidris fuscicollis, Pluvialis squatarola and Tringa melanoleuca, used foraging paths that tended to maximize the distance from burrows, especially the distance to larger burrows. A field experiment showed that this was not necessarily due to a decrease in the availability of polychaetes near the crab burrows. A combination of landscape measurements and satellite images showed that crab beds covered up to 40% of the intertidal area of the Mar Chiquita coastal lagoon (37°40′S, Argentina), and nearly 100% of the intertidal area of the Bahia Blanca estuary (38°48′-39°25′S, Argentina). These two estuaries are located along the migratory flyway of Neotropical migratory shorebirds, but the Bahia Blanca estuary (area∼110,000 ha) shows a much lower shorebird diversity than Mar Chiquita (area∼4500 ha). The most common species in Bahia Blanca is the two-banded plover C. falklandicus, the species least affected by crabs at Mar Chiquita and which prefers to use high-density crab areas as foraging sites. The oystercatcher Haematopus palliatus was also most abundant in high-density crab areas, but they used these areas for resting. The abundances of preys varied during the study period and between the crab density areas, indicating that the use of these areas by birds is independent of crab density. However, burrowing crabs affect the depth distribution of polychaete and thus their availability to shorebirds. We suggest that this shorebirds-burrowing organism interaction could be generalized for other intertidal estuarine habitats.     

Habitat use and early winter movements by juvenile Atlantic cod in a coastal area of Newfoundland

Acoustic telemetry was used to monitor the movements and landscape scale habitat use of age 2–3 year juvenile Atlantic cod Gadus morhua in Newman Sound, a coastal fjord of Newfoundland, during late autumn and early winter (24 October 1999 to 22 January 2000). Substratum, bathymetric relief and depth use were studied to determine if it differed from the pattern expected given an absence of selectivity (habitat use proportional to habitat availability). Prior to winter migrations, most Atlantic cod maintained small home ranges (0·5–33·4 ha, median = 2·1 ha) although a few individuals moved more widely. No relationship between total length (L_T) and home range size was detected. In Inner Newman Sound, Atlantic cod occupied depths of 10–29 m more than expected given availability, while depths of 0–9 and 40–59 m were underutilized. No significant relationship between depth and L_T was detected. Areas of medium (5–10%) or high (>10%) bathymetric relief and boulder or kelp habitats were used significantly more than expected given the availability of these habitats. Sand and eelgrass substrata were underutilized given availability, although many Atlantic cod used sandy‐bottomed areas to some degree. Flexibility in habitat use by the individuals that were studied suggested reduced predation risk relative to younger conspecifics. Winter migrations to deeper water beyond Newman Sound began in mid‐November, coinciding with the disappearance of the thermocline, and continued until 27 December. Approximately 30% of monitored individuals did not migrate and maintained their home ranges into the winter season. No significant differences in L_T between migrating and resident groups were detected, however, the condition of migrating fish was significantly higher than in resident fish. This finding supports the theory that feeding history plays a role in the decision to undertake migration.     

Effects of native predators and eelgrass habitat structure on the introduced Asian mussel Musculista senhousia (Benson in Cantor) in southern California

The ability of predators to control the abundance of non-native species has been little explored in marine systems. Native predators may be used to control non-native species or may confer invasion resistance to communities if predation rates on invaders are density-dependent. We studied the response of southern California native predators to the density of Musculista senhousia (Benson in Cantor, 1842), a small, fast growing mussel that has been introduced from Japan to several coastlines worldwide. We performed field experiments to determine if M. senhousia proportional mortality is density-dependent and if eelgrass Zostera marina L. habitat structure influenced mussel density-dependent mortality. We also evaluated the effect of seagrass habitat structure on the aggregative and functional responses of the predatory gastropod Pteropurpura festiva (Hinds, 1844) to Asian mussel density. In the summer of 2002, P. festiva aggregated in plots with high mussel density and was responsible for nearly all predation on M. senhousia. However, M. senhousia proportional mortality was inversely density-dependent at all levels of eelgrass above-ground and below-ground habitat structure. Asian mussel proportional mortality also was inversely density-dependent and was not influenced by eelgrass habitat structure in the spring of 2004 when wading birds were the chief predator of mussels. In contrast to results for mussel proportional mortality, the aggregative and functional responses of P. festiva varied with seagrass habitat structure. P. festiva density increased with Asian mussel density in plots with low simulated habitat structure, but the relationship between P. festiva density and Asian mussel density was parabolic at zero, intermediate and high levels of habitat structure. In field enclosures, P. festiva exhibited a Type I (linear) functional response to Asian mussel density at low levels of eelgrass structure, and a Type II (hyperbolic) functional response to mussel density at high levels of eelgrass structure. Our results and those of others suggest that the degree to which local benthic communities in southern California are resistant to Asian mussel invasion depends on habitat structure, mussel settlement rates, and the density and diversity of predators.     

Habitat selection of juvenile banana prawns, Penaeus merguiensis de Man: Testing the roles of habitat structure, predators, light phase and prawn size

The effects of fish predators, light phase, habitat structure and prawn size on the habitat preferences of juvenile Penaeus (Fenneropenaeus) merguiensis de Man were examined with laboratory experiments. The behaviour of juvenile P. merguiensis within habitats of different structural complexity was also examined. Experiments were carried out in a tank (1.8 m diameter) divided into four habitats representing: bare substratum, leaf litter (little vertical structure), mangrove pneumatophores (regular vertical structure) and mangrove woody debris (heterogeneous vertical structure). The location of 10 prawns was monitored over 270 min (135 min light and 135 min dark), with different prawns five times for each combination of prawn size class, and predator (no predator; Arius graeffei Kner and Steindachner and Lates calcarifer Bloch).In the absence of predators and during the light phase, when observations on prawn behaviour were made, swimming was the most common behaviour (of seven mutually exclusive behavioural categories) with few differences in behaviour between sizes. All size classes of juvenile P. merguiensis selected vertical structure (mangrove debris and pneumatophores) over low vertical structure (leaf litter and bare substratum), in both light and dark conditions and in the presence or absence of predators. When L. calcarifer was present, the selection by prawns of the mangrove-debris habitat increased significantly. This was attributed to an increase in predation risk in the other habitats. L. calcarifer rarely pursued prey amongst the mangrove-debris structure, compared to habitats with less heterogeneous vertical structure (pneumatophores, leaf litter and bare substratum).     

Behavioural mechanisms of sea stars (Asterias vulgaris Verrill and Leptasterias polaris Müller) and crabs (Cancer irroratus Say and Hyas araneus Linnaeus) preying on juvenile sea scallops (Placopecten magellanicus (Gmelin)), and procedural effects of scallop tethering

We compared predation rates and behaviours of sea stars (Asterias vulgaris and Leptasterias polaris) and crabs (Cancer irroratus and Hyas araneus) preying on juvenile sea scallops (Placopecten magellanicus, 25-35 mm shell height) in laboratory. These predatory species co-occur with sea scallops on the sea bed of the Gulf of St. Lawrence, Canada, and limit scallop survival in seeding operations. We also examined, under controlled conditions, the effect of tethering scallops on predator-prey interactions. Predation rates, time budgets and encounter behaviours observed for A. vulgaris and C. irroratus preying on free (untethered) scallops were comparable to previous studies. C. irroratus were more effective predators as they consumed 3.1 scallops predator^− 1 day^− 1, although they spent only 0.9% of their time searching for prey. A. vulgaris consumed 0.9 scallops predator^− 1 day^− 1 and spent 7.6% of their time searching. Sea stars L. polaris had a lower predation rate (0.02 scallop predator^− 1 day^− 1) than A. vulgaris. The frequent avoidance behaviour of L. polaris and its low ability to capture scallops support the notion that scallops are not a main component of this sea star's diet. Crabs H. araneus had similar predation rates (1.3 scallops predator^− 1 day^− 1) and behaviours to C. irroratus, although the probability of consumption upon capture was affected by relatively high numbers of rejections and post-capture escapes of scallops. As expected, the tethering procedure increased predation rate of L. polaris (about 19 times higher), but surprisingly did not significantly affect that of A. vulgaris. Examination of behaviours indicated that A. vulgaris offered tethered scallops tended to have a higher probability of capture, but spent less time searching for prey (possibly because satiation was reached) than A. vulgaris offered free scallops. Predation rates and behaviours of both crab species were not affected by tethering, since encounter rate was the primary determinant of crab-scallop interactions. Identification and quantification of behaviours underlying the predation process allowed us to mathematically model predator-related mortality for the four predator species.