Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Tissue distribution and excretion of 99mTc-disofenin in three marine species: Pleuronectes americanus (winter flounder), Homarus americanus (lobster), and Mya arenaria (soft-shell clam)

To determine the pharmacokinetics of a small lipophilic molecule in vivo, the distribution and accumulation of ^99mTc-radiolabelled disofenin (diisopropylacetanilide iminodiacetic acid) were traced during 1991–1992 by scintigraphy and gamma well counting in winter flounder (Pleuronectes americanus collected from Boston Harbor and Long Island Sound in 1992), lobsters (Homarus americanus collected from Massachusetts Bay in 1991), and soft-shell clams (Mya arenaria purchased in 1991). The agent was distributed throughout the bodies of lobsters within 12 s, throughout flounder within 40 s, and throughout clams within 2 min. It was concentrated most strongly by the liver of flounder, which contained 61.2±7.8% of the injected dose within 1 h of injection, and by the lobster hepatopancreas. Accumulation also occurred in the flounder kidney, lobster antennal glands, and the kidney and pericardial glands of clams. The compound was rapidly excreted from the flounder liver into the gall bladder, and from the lobster hepatopancreas into the stomach. The data suggest its excretion from the lobster antennal glands and clam kidneys. The rate of clearance of disofenin from the body varied among species: 99±2.1% of the initial dose remained in flounder sampled 16 to 24 h after injection, compared to 80.5±7% remaining in the lobster after 15 h, and 87.4±5.9% remaining in clams after 27 h. The clearance rates in flounder and lobster are considered to be minimum values because of the lack of gut activity in unfed individuals. Overall, these in vivo tracer studies establish the utility of scintigraphy for assessing the uptake and excretion of a lipid soluble compound in different taxa, and may be applicable for evaluating disease and/or environmental effects on organ function in marine animals.     

Long-term trends in invertebrate–habitat relationships under protected and fished conditions

Few studies examine the long-term effects of changing predator size and abundance on the habitat associations of resident organisms despite that this knowledge is critical to understand the ecosystem effects of fishing. Marine reserves offer the opportunity to determine ecosystem-level effects of manipulated predator densities, while parallel monitoring of adjacent fished areas allows separating these effects from regional-scale change. Relationships between two measures of benthic habitat structure (reef architecture and topographic complexity) and key invertebrate species were followed over 17 years at fished and protected subtidal rocky reefs associated with two southern Australian marine reserves. Two commercially harvested species, the southern rock lobster (Jasus edwardsii) and blacklip abalone (Haliotis rubra) were initially weakly associated with habitat structure across all fished and protected sites. The strength of association with habitat for both species increased markedly at protected sites 2 years after marine reserve declaration, and then gradually weakened over subsequent years. The increasing size of rock lobster within reserves apparently reduced their dependency on reef shelters as refuges from predation. Rising predation by fish and rock lobster in the reserves corresponded with weakening invertebrate–habitat relationships for H. rubra and sea urchins (Heliocidaris erythrogramma). These results emphasise that animal–habitat relationships are not necessarily stable through time and highlight the value of marine reserves as reference sites. Our work shows that fishery closures to enhance populations of commercially important and keystone species should be in areas with a range of habitat features to accommodate shifting ecological requirements with ontogenesis.     

Mitochondrial DNA variation in the European lobster (<Emphasis Type="Italic">Homarus gammarus</Emphasis>) throughout the range

The genetic differentiation of the European lobster (Homarus gammarus) was investigated in 3,283 individuals from 44 population samples throughout its geographical distribution (Norway to Greece) by means of polymerase chain reaction restriction fragment length polymorphism analysis of a 3-kb mitochondrial DNA segment. Ninety composite haplotypes were revealed with the number of haplotypes in each population sample ranging from 4 to 31. Private haplotypes were found at very low frequencies. The global exact test of sample differentiation based on composite haplotype frequencies was statistically significant. F_ST analyses also showed significant heterogeneity among the European lobster population samples (F_ST=0.078, P<0.001). This differentiation was mainly due to the population samples from northern Norway, the Netherlands, and the Mediterranean Sea. These samples differentiated from the rest due to reduced gene diversity rather than to unique haplotypes. The relationships among these samples were illustrated with cluster analyses; four major distinct groups were evident: Mediterranean, northern Norway, Netherlands, and the remaining Atlantic samples. Based on the low degree of differentiation revealed in the European lobster and its limited capacity for dispersal, the most probable hypothesis is that all populations have been established from a common refuge after the end of the last Ice Age, that is, within the past 15,000 years. The results of this study show that mitochondrial DNA is a powerful tool for the determination of the genetic structure among lobster samples, which is important for a proper management policy designed to protect and to cultivate this species.Communicated by O. Kinne, Oldendorf/Luhe     

Succession in sea urchin and seaweed abundance in Nova Scotia,Canada

From 1980 through 1983 sea urchin mass mortalities were at least 245 000 t. The habitat released to seaweed is expected to support a standing crop of 1.8 million tons and an annual production of 7 million tons. An extensive survey of 2 900 km of Nova Scotia shoreline revealed that 511 km² of habitat (rock bottom less than 15 m deep) was available to seaweeds. Areas most exposed to ocean swells bordered 26% of the shoreline length but included 82% of the seaweed habitat. Before mass mortalities, seaweed refuges from sea urchin grazing included very sheltered habitats, very exposed habitats, and boulders set among shifting sand; however, these were less than 10% of the total habitat. In the 1 to 2 years following mass mortalities seaweed cover and occurrence increased significantly for all of five categories of wave exposure and for six of seven algal taxa respectively. In at least three locations the sea urchin population began to recover as a result of larval recruitment but in 1983 was again reduced by disease. Sea urchins appear to lack natural resistance to disease, at least at warm temperatures. In the laboratory all sea urchins collected from the Bay of Fundy, southern Nova Scotia, eastern Newfoundland, and an area recovering from mass mortalities died following exposure to diseased animals. Also, field mortalities included a wide variety of habitats. Interviews of Nova Scotia lobster fishermen revealed that sea urchin mass mortalities have probably occurred before in this century, but infrequently.     

Shelter selection of the spiny lobster <Emphasis Type="Italic">Palinurus elephas</Emphasis> under different levels of <Emphasis Type="Italic">Octopus vulgaris</Emphasis> predation threat

The skill of recognizing and reacting to predators is often based on a learned component. Few studies have examined the role of learning in spiny lobster anti-predator behavior. We investigated whether European spiny lobster (Palinurus elephas) shelter selection is influenced by olfactory stimuli released by one of the most common lobster predators, the common octopus (Octopus vulgaris), and whether the behavioral response to octopus chemical stimuli is innate or influenced by experience. In experimental arenas, we conditioned wild-caught lobsters with three levels of predation threat: no threat, with no predator–prey interaction; medium threat, with odor and visual predator cues only; high threat, active predation risk. We subsequently tested the shelter choice of the conditioned lobster under different experimental conditions: (1) shelter plus seawater; (2) shelter plus seawater plus chemical octopus cue. Our results showed significant differences in mean shelter occupancy with conditioning level. We conclude that P. elephas individuals use chemosensory systems in predator-avoidance mechanisms. Moreover, lobsters subject to a training period of high-level predation threat were able to learn the octopus chemical stimuli and treat its odor as a cue related to predation risk. The findings relative to the spiny lobster learning abilities could be an important tool for future management of lobster populations, e.g., by re-introduction of reared juveniles, which have not yet experienced predation.     

Circadian shelter occupancy patterns and predator–prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon

The spiny lobster, Panulirus argus, is predominantly nocturnal, remaining inside shelters during the day and foraging outside at night, presumably to minimize predation risk. Predation risk generally decreases with increasing lobster size. Therefore, this study examined the hypothesis that size would influence this basic circadian pattern. Video cameras continuously recorded the shelter occupancy of juvenile lobsters (n = 72) having a carapace length (CL) of 30–62 mm that were tethered to shelters in a shallow reef lagoon. The lobsters’ shelter occupancy was 100% during the day, but declined linearly from shortly before sunset to a minimum of 50% shortly after midnight and then increased linearly, reaching 100% by 1 h after sunrise. The percent time the lobsters spent in the shelters followed a similar trend, but there was wide variability at night (0–100%) for individual lobsters. Lobsters left their shelters 2–30 times night⁻¹, with a majority of excursions lasting <10 min. These results suggest that juvenile P. argus minimize predation risk by remaining in their shelters as long as possible but offset the energetic cost of this behavior by foraging close to their shelters for several short periods at night. This emergence pattern contrasts with those of early benthic phase lobsters (<15 mm CL), which seldom leave their shelters, and adults (>80 mm CL), which have a dusk/early evening peak in activity and leave the shelter for extended periods of time during the night. Furthermore, a minimum shelter occupancy in the middle of the night appears especially well adapted to avoid exposure to daytime predators. Videotaped observations also included interactions between lobsters and two dominant lobster predators, the triggerfish, Balistes capriscus, and the octopus Octopus cf. vulgaris. Lobsters responded differently to these predators: remaining in the shelter when attacked by a triggerfish and fleeing the shelter when attacked by an octopus. Triggerfish were nearly twice as likely to attack a lobster that was outside of the shelter than inside. Once under attack, however, a lobster had nearly the same chance of surviving if it was inside or outside. Results suggest that the patterns of shelter use and emergence change as lobsters grow, probably reflecting the interplay between perception of predation risk and the need to forage. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Recovery of corals a decade after a bleaching event in Dubai, United Arab Emirates

Elevated sea surface temperatures in the late 1990s were associated with widespread coral mortality in the Arabian Gulf, particularly in Acropora dominated areas. This study investigates the composition, condition, and recruitment patterns of coral communities in Saih Al-Shaib, Dubai, United Arab Emirates, a decade after mass bleaching. Five statistically distinct communities were identified by cluster analysis, with grouping optimized from 17 significant indicator species. Overall, 25 species of scleractinian coral were observed, representing 35 ± 1.6% coral cover. Densities of recruits were low (0.8 ± 0.2 m⁻²), and composition generally reflected that of the surrounding adult community. Ten years after mass mortality, Acropora dominated assemblages were observed in three of the six sites examined and coral cover (41.9 ± 2.5%) was double post-bleaching cover. One shallow near-shore site appears to have had recovery of Acropora reset by a further bleaching event in 2002. However, the prevalence of young Acropora colonies here indicates that recovery may recur in several years. One area formerly dominated by Acropora is now dominated by faviids and poritids, with adult and juvenile composition suggesting this dominance shift is likely to persist. Porites lutea and Porites harrisoni dominated communities were negligibly impacted by the bleaching events, and the limited change in coral cover and composition in intervening years likely results from slow growth and low recruitment. Despite strong recovery of several dominant Acropora species, five formerly common species from this area were not observed suggesting local extinction. Dubai coral communities exhibit both resistance and resilience to elevated sea temperatures. The conservation of these patch reefs is warranted given the predicted increase in bleaching events, and the role that these communities may play in regional recovery.     

Creation of a Gilded Trap by the High Economic Value of the Maine Lobster Fishery

Abstract: Unsustainable fishing simplifies food chains and, as with aquaculture, can result in reliance on a few economically valuable species. This lack of diversity may increase risks of ecological and economic disruptions. Centuries of intense fishing have extirpated most apex predators in the Gulf of Maine (United States and Canada), effectively creating an American lobster (Homarus americanus) monoculture. Over the past 20 years, the economic diversity of marine resources harvested in Maine has declined by almost 70%. Today, over 80% of the value of Maine's fish and seafood landings is from highly abundant lobsters. Inflation‐corrected income from lobsters in Maine has steadily increased by nearly 400% since 1985. Fisheries managers, policy makers, and fishers view this as a success. However, such lucrative monocultures increase the social and ecological consequences of future declines in lobsters. In southern New England, disease and stresses related to increases in ocean temperature resulted in more than a 70% decline in lobster abundance, prompting managers to propose closing that fishery. A similar collapse in Maine could fundamentally disrupt the social and economic foundation of its coast. We suggest the current success of Maine's lobster fishery is a gilded trap. Gilded traps are a type of social trap in which collective actions resulting from economically attractive opportunities outweigh concerns over associated social and ecological risks or consequences. Large financial gain creates a strong reinforcing feedback that deepens the trap. Avoiding or escaping gilded traps requires managing for increased biological and economic diversity. This is difficult to do prior to a crisis while financial incentives for maintaining the status quo are large. The long‐term challenge is to shift fisheries management away from single species toward integrated social‐ecological approaches that diversify local ecosystems, societies, and economies.     

Growth of the rock lobster Jasus lalandii and its relationship to benthos

Growth increments of the male rock lobster Jasus lalandii vary systematically with location within a small (38 km²) fishing ground off the South African west coast. High growth rates were recorded from an area where the availability of benthos suitable as rock-lobster food was also high. Increments could also be shown to vary significantly in different years, but over the size range examined, growth did not appear to be a function of size. Benthic biomass was found to be inversely related to depth. However, comparisons between biomass values from the same depths in two different areas of the fishing ground suggested that biomass was higher in areas which were in proximity to shallow water, where extensive kelp beds were found. Mussels (Aulacomya magellanica) formed an important constituent of the rock lobster's diet; these were prevalent at depths between 20 and 40 m, and especially in areas where rock-lobster growth rates were high.     

A whole-animal bioassay for the determination of the food attractants of the lobster Homarus gammarus

A semi-quantitative assay is described for assessing the effectiveness of bait extracts for the lobster Homarus gammarus (L.), using an extract of the squid Loligo vulgaris Lamarck as a standard. Fractionation of this extract indicated that, while no single component was as attractive to lobsters as the whole, the recombination of the various components yielded a solution which was as attractive as the original extract. It appears likely that several different classes of chemosensory cell must be simultaneously stimulated for the initiation of feeding behaviour in the lobster.     

Acceptability and digestion of diets fed to larval stages ofHomarus gammarus and the role of dietary conditioning behaviour

Whilst the role of chemoreception in juvenile crustacean feeding behaviour has received considerable attention, the chemosensory responses of larval planktonic crustacean stages have been neglected. The present study investigated the acceptability of processed, microparticulate and microencapsulated diets and the possible role of chemoattractants for larval and juvenile stages ofHomarus gammarus (Linnaeus) and juvenile stages ofPalaemon elegans (Rathke). Larval and juvenile lobster were obtained from MAFF (Ministry of Agriculture, Fisheries and Food, Conwy, UK), and juvenileP. elegans were collected from the Menai Straits (Gwynedd, UK) during the summer months of 1985/1986. During the larval stages, food capture results from chance encounter, and the role of low molecular-weight chemoattractants appears to be in the determination of the edible from the inedible. Dietary conditioning behaviour in decapod larvae, which has until now not been investigated, enables the larvae to discriminate more rapidly between particles of different digestibilities. The digestibility and assimilation efficiency displayed on artificial diets in the present study suggest that lobster larvae possess limited enzymatic capabilities, although later stages are more efficient and display longer food-retention time. Growth (0.114 mm d^–1) and survival (80.0%) of lobster larvae fed natural diets were significantly better (F=14.8;P=0.001, andF=54.0,P=0.001, respectively) than of individuals fed artificial diets. Although pre-digested artificial diets (ingredients pre-treated with pancreatin) resulted in higher survival (74 and 63%) compared to microencapsulated diets (37%), there was no significant growth difference between the artificial diets tested.     

Seasonal and long-term variations in the body size of planktonic copepods in the northern Baltic Sea

Seasonal and long-term variations in the body size of planktonic copepods were studied in the northern Baltic Sea. The influence of temperature, salinity and phytoplankton concentration to the body size of Eurytemora affinis, Acartia bifilosa and Limnocalanus macrurus (Calanoida) was examined at three sea areas, differing in their hydrographical and trophic conditions (an archipelago area and an enclosed bay on the SW coast of Finland in 1992 and at an open sea station in the Gulf of Bothnia in 1991). There was an inverse relationship between copepod body length and temperature, while there was a direct effect of phytoplankton concentration. According to multiple regression analysis, the relative importance of these factors varied according to species, developmental stage and study area. In the archipelago area, copepod body size was mainly determined by temperature, while in the open Gulf of Bothnia, phytoplankton concentration was usually more important. Interannual variation in the summer body length of E. affinis and A. bifilosa was examined using samples collected over 18 years (1967 to 1984) in the archipelago area. In this analysis, the inverse relationship between water temperature and copepod body size disappeared; in E. affinis the average female length correlated positively with summer temperature. We suggest that the length and timing of the study period greatly influence the investigator's view of the factors regulating copepod body size.     

Between-individual variation in haemocyanin concentrations in the Norway lobster<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> following exposure to hypoxia and manganese

Environmental hypoxia and the release of reduced manganese (Mn) are often combined stress factors in marine sediments. Previous investigations have shown that hypoxia induces variable responses in the concentration of the respiratory pigment haemocyanin (Hc) of crustaceans. A recent study demonstrated that the magnitude and direction of the changes was dependent on the initial level of Hc in individuals of the Norway lobster Nephrops norvegicus. The changes also took place within 24 h rather than days as suggested by previous studies. In this present study we investigated changes of Hc in individual N. norvegicus in uncontaminated sea water after exposure to and release from realistic hypoxic stress (PO ₂=6.0 kPa). Furthermore we investigated how concomitant exposure to realistic concentrations of manganese (20 mg l ^-1) modified the responses we observed. We confirmed a between-individual variation in response and a change in Hc towards an optimum level after hypoxia, but also showed that changes that took place post-hypoxic exposure occurred over a timescale similar to that of the hypoxic response itself. Manganese exposure resulted in no significant changes in Hc whereas Mn exposure combined with hypoxia resulted in a significant decrease (15%) in all individuals independent of initial Hc. N. norvegicus was thus unable to compensate for the effects of hypoxia by synthesis of Hc after exposure to naturally occurring Mn concentrations.     

Short-term effects of an oil spill on the West coast of the Cape Peninsula, South Africa

In 1994, the sinking of the ‘Apollo Sea’ off the West coast of South Africa led to the deposition of ca. 2 500 tons of heavy fuel oil over 150 km of coastline. The impact of the spill on rocky shore invertebrates, rock-pool fish fauna and rock lobsters was assessed by conducting surveys shortly after the spill, and again two months later. Where possible, results of these surveys were compared with existing data from before the sinking of the Apollo Sea. Among the fish fauna of rock-pools, changes in total density of fishes were largely due to changes in the abundance ofClinus superciliosus and were within the range of natural variation for the species and the community as a whole. Community structure of the rock-pool fish fauna also remained unaffected. At three of four impacted rocky shore sites no changes could be detected in overall benthic community structure, although the winkleNodilittorina africana was affected. At the fourth site, a boulder beach, statistical analysis showed distinct differences in community structure between heavily and lightly contaminated areas, as well as between all areas compared with previously existing data. There were also significant changes in the lightly oiled areas between the first and second surveys after the spill. Oil-fouled lobster were found at one of three sites investigated. Ca. 7 % of the seabed in this particular area was polluted. Antennae and forelegs of almost all lobsters in the vicinity of the oil were fouled. Examination of the gut-contents confirmed that oil inhibits the ability of lobster to feed. Although there was no evidence that mortalities of lobster were caused by the spill, growth rates may be reduced by the decreased feeding rate. Overall, the impact of the spill was strikingly less than might have been predicted from the effects of other oil spills. Nomenclature: Names of fish species follow Smith & Heemstra (1986).     

Locomotory behaviour and swimming performance of the Norway lobster,Nephrops norvegicus,in the presence of an acoustic tag

Observations have been made on the locomotory behaviour and swimming performances of the Norway lobster, Nephrops norvegicus (L.), fitted with an acoustic transmitter ventrally under the cephalothorax. The walking behaviour of adult males (44 mm carapace length) appeared to be unaffected, but the tag caused significant reductions in certain measures of tail-flip swimming performance such as swimming speed and endurance. Flume-tank experiments in low water currents suggested that the transmitter would increase hydrodynamic drag during swimming by 9 to 32%, depending on lobster size. Given the weight and dimensions of the acoustic transmitters currently available, it is considered advisable to confine acoustic tracking studies to relatively large N. norvegicus.Correspondence to: C. J. Chapman     

Modeling an exploited rocky coastal ecosystem: Bahia Tortugas, Mexico

A trophic structure model of the rocky coastal ecosystem in Bahia Tortugas, Mexico was constructed using Ecopath software to represent the main biomass flows in the system. Data for the model came from field observations (biomass estimates, stomach contents, and ecological observations for sea snails, abalones, lobster, some demersal finfishes, and macroalgae) carried out through ten field trips from 2006 to 2008. The results provide a snapshot of how the ecosystem operates. The model considers 23 functional groups. The total system throughput was 553 t/km²/year, 57% corresponds to internal consumption, 28% to respiration, 14% becomes detritus, and only 1% is removed through commercial fishing. The model suggests that even for exploited populations, predation and competition are heavier stresses than current fishing effort; however, because spiny lobster showed the second highest keystoneness’ index value, increasing fishing pressure on this group could strongly impact the entire ecosystem. We believe that this model has the potential to support management by allowing the exploration of the potential impacts of different fishing decisions at ecosystem level.     

Biochemical composition of a meso-bathyal lobster

To characterise some traits of the life strategies of Polycheles typhlops, a deep-sea small lobster usually encountered as by-catch within deep trawls in the Southern Tyrrhenian Sea (Mediterranean Sea), we analysed the biochemical composition of the caudal muscles of 49 specimens collected in March and April 2007. Differences in the biochemical composition between females and males were weak, but relevant differences in protein (>90% of the organic C) and lipid (4–6%) contents were observed between developmental stages both in females and males. These differences are likely to be related to different food items and/or to the metabolic shifts associated with their reproduction. We argue that this deep-sea lobster could cover a relevant role in the meso-pelagic food webs and attract future commercial interest.     

Cryptic species within the commercially most important lobster in the tropical Atlantic, the spiny lobster Panulirus argus

Panulirus argus (Latreille in Ann Mus Hist Nat Paris 3:388–395, 1804) is the lobster of greatest economic importance throughout its distribution. In this study, mitochondrial (Cytochrome Oxidase I and 16S ribosomal genes) and nuclear (Adenine Nucleotide Transporter gene) sequences were used to evaluate the taxonomic status of P. argus sampled from five sites in the Caribbean Sea and nine sites in the Southwest Atlantic. Phylogenetic analyses indicate that lobsters from the two regions form two monophyletic groups with a molecular divergence similar to that observed between distinct congeneric lobster species and much larger than that found between conspecific lobster populations. Therefore, the Caribbean and the Southwest Atlantic lobster populations originally attributed to P. argus belong to different species, with an estimated time of isolation of around 16 Million years. An important consequence of these findings is that the fisheries of spiny lobsters from the Caribbean and the Southwest Atlantic species must be managed separately.