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1. Connectivity of marine populations and ecosystems is crucial to maintaining and enhancing their structure, distribution, persistence, resilience and productivity. Artificial hard substrate, such as that associated with oil and gas platforms, provides settlement opportunities for species adapted to hard substrates in areas of soft sediment. The contribution of artificial hard substrate and the consequences of its removal (e.g. through decommissioning) to marine connectivity is not clear, yet such information is vital to inform marine spatial planning and future policy decisions on the use and protection of marine resources. 2. This study demonstrates the application of a social network analysis approach to quantify and describe the ecological connectivity, informed by particle tracking model outputs, of hard substrate marine communities in the North Sea. Through comparison of networks with and without artificial hard substrate, and based on hypothetical decommissioning scenarios, this study provides insight into the contribution of artificial hard substrate, and the consequence of decommissioning, to the structure and function of marine community connectivity. 3. This study highlights that artificial hard substrate, despite providing only a small proportion of the total area of hard substrate, increases the geographic extent and connectivity of the hard substrate network, bridging gaps, thereby providing ‘stepping stones’ between otherwise disconnected areas of natural hard substrate. Compared to the baseline scenario, a decommissioning scenario with full removal of oil and gas platforms results in a nearly 60% reduction in connectivity. Such reduction in connectivity may have negative implications for species’ distribution, gene flow and resilience following disturbance or exploitation of marine hard substrate communities. 4. Synthesis and applications. Social network analysis can provide valuable insight into connectivity between marine communities and enable the evaluation of impacts associated with changes to the marine environment. Providing standardized, transparent and robust outputs, such a tool is useful to facilitate understanding across different disciplines, including marine science, marine spatial planning and marine policy. Social network analysis therefore has great potential to address current knowledge gaps with respect to marine connectivity and crucially facilitate assessment of the impacts of changes in offshore substrate as part of the marine spatial planning process, thereby informing policy and marine management decisions.

As the EU's commitment to renewable energy is projected to grow to 20% of energy generation by 2020, the use of marine renewable energy from wind, wave and tidal resources is increasing. This literature review (233 studies) (i) summarizes knowledge on how marine renewable energy devices affect benthic environments, (ii) explains how these effects could alter ecosystem processes that support major ecosystem services and (iii) provides an approach to determine urgent research needs. Conceptual diagrams were set up to structure hypothesized cause-effect relationships (i.e. paths). Paths were scored for (i) temporal and spatial scale of the effect, (ii) benthic sensitivity to these effects, (iii) the effect consistency and iv) scoring confidence, and consecutively ranked. This approach identified prominent knowledge gaps and research needs about (a) hydrodynamic changes possibly resulting in altered primary production with potential consequences for filter feeders, (b) the introduction and range expansion of non-native species (through stepping stone effects) and, (c) noise and vibration effects on benthic organisms. Our results further provide evidence that benthic sensitivity to offshore renewable effects is higher than previously indicated. Knowledge on changes of ecological functioning through cascading effects is limited and requires distinct hypothesis-driven research combined with integrative ecological modelling.

Non-consumptive effects of predators can have important impacts on aquatic food webs, but there are few data on how predators change the behaviour of Caribbean reef fishes. Such changes may include behavioural responses to the invasive predatory lionfish (Pterois volitans/P. miles). This study used an aquarium experiment to examine the behaviour of herbivorous parrotfish (Scarus iseri) in the absence of other fish (control), with a non-piscivore present, and with a predatory threat from a native grouper or lionfish. Treatments were repeated with and without additional parrotfish shelters to examine the potential effects of degraded reefs (loss of refuges). Using video, parrotfish behaviours (sheltering, swimming in open areas, foraging, aggressive conspecific interactions, bite rates, and shoaling behaviour) were recorded for groups of four parrotfish. Compared to the control, the average number of parrotfish hiding was reduced by 65 % and foraging shoals were 10 % larger when threatened by grouper, likely as a specific response to an ambush predator. When exposed to lionfish, parrotfish reduced their bite rates by 50 %, possibly to be more vigilant of this predator’s unique stalking behaviour. The absence of additional shelter had limited effects although parrotfish formed 10 % larger shoals when swimming in open water, potentially as a defensive behaviour because of a perceived lack of refuges. The reduction in parrotfish bite rates caused by lionfish may have important demographic consequences. Furthermore, parrotfishes are important grazers of macroalgae, and these behavioural changes may exacerbate the direct effects of lionfish predation and potentially affect reef benthic dynamics.

© The Ecological Society of America The removal of thousands of structures associated with oil and gas development from the world's oceans is well underway, yet the environmental impacts of this decommissioning practice remain unknown. Similar impacts will be associated with the eventual removal of offshore wind turbines. We conducted a global survey of environmental experts to guide best decommissioning practices in the North Sea, a region with a substantial removal burden. In contrast to current regulations, 94.7% of experts (36 out of 38) agreed that a more flexible case-by-case approach to decommissioning could benefit the North Sea environment. Partial removal options were considered to deliver better environmental outcomes than complete removal for platforms, but both approaches were equally supported for wind turbines. Key considerations identified for decommissioning were biodiversity enhancement, provision of reef habitat, and protection from bottom trawling, all of which are negatively affected by complete removal. We provide recommendations to guide the revision of offshore decommissioning policy, including a temporary suspension of obligatory removal.

Predator avoidance is fundamental for survival and it can be particularly challenging for prey animals if physical movement away from a predatory threat is restricted. Many sharks and rays begin life within an egg capsule that is attached to the sea bed. The vulnerability of this sedentary life stage is exacerbated in skates (Rajidae) as the compulsory ventilatory activity of embryos makes them conspicuous to potential predators. Embryos can reduce this risk by mediating ventilatory activity if they detect the presence of a predator using an acute electrosense. To determine how early in embryonic life predator elicited behavioral responses can occur, the reactions of three different age groups (1/3 developed, 2/3 developed, and near hatching) of embryonic thornback rays Raja clavata were tested using predator-type electric field stimuli. Egg capsules were exposed to continuous or intermittent stimuli in order to assess varying predator-type encounter scenarios on the ventilatory behavior of different developmental stages. All embryos reacted with a "freeze response" following initial electric field (E-field) exposure, ceasing ventilatory behavior in response to predator presence, demonstrating electroreceptive functionality for the first time at the earliest possible stage in ontogeny. This ability coincided with the onset of egg ventilatory behavior and may represent an effective means to enhance survival. A continuous application of stimuli over time revealed that embryos can adapt their behavior and resume normal activity, whereas when presented intermittently, the E-field resulted in a significant reduction in overall ventilatory activity across all ages. Recovery from stimuli was significantly quicker in older embryos, potentially indicative of the trade-off between avoiding predation and adequate respiration. © 2015 Wiley Periodicals, Inc. Develop Neurobiol 76: 721-729, 2016.

Introduction Deforestation significantly impacts large carnivores that depend on large tracts of interconnected forest habitat and that are sensitive to human activities. Understanding the relationship between habitat use and spatial distribution of such species across human modified landscapes is critical when planning effective conservation strategies. This study assessed the presence of potential landscape connectivity thresholds resulting from habitat fragmentation associated with different deforestation patterns using a scale-based approach that links species-specific home ranges with the extent of anthropogenic activities. The objectives were (1) to quantify the spatial and temporal distribution of natural vegetation for five common deforestation patterns and (2) to evaluate the connectivity associated with these patterns and the existence of potential thresholds affecting jaguar dispersal. The Bolivian lowlands, located within jaguar conservation units, were analysed with landscape metrics to capture the spatial and temporal changes within deforested areas and to determine potential impact on jaguar connectivity and connectivity thresholds for dispersal. Results Over the period of 1976–2005, the amount of natural vegetation has decreased by more than 40% in all locations with the biggest changes occurring between 1991 and 2000. Landscape spatial structure around jaguar locations showed that jaguars used areas with mean proportion of natural areas = 83.14% (SE = 3.72%), mean patch density = 1.16 patches/100 ha (SE = 0.28 patches/100 ha), mean patch area = 616.95 ha (SE = 172.89 ha) and mean edge density = 705.27 m/ha (SE = 182.19 m/ha).We observed strong fragmentation processes in all study locations, which has resulted in the connectivity of jaguar habitat decreasing to

The considerable extent of construction and operation of marine renewable energy developments (MRED) within U.K. and adjacent waters will lead, among other things, to the emission of electromagnetic fields (EMF) and subsea sounds into the marine environment. Migratory fishes that respond to natural environmental cues, such as the Earth's geomagnetic field or underwater sounds, move through the same waters that the MRED occupy, thereby raising the question of whether there are any effects of MRED on migratory fishes. Diadromous species, such as the Salmonidae and Anguillidae, which undertake large-scale migrations through coastal and offshore waters, are already significantly affected by other human activities leading to national and international conservation efforts to manage any existing threats and to minimize future concerns, including the potential effect of MRED. Here, the current state of knowledge with regard to the potential for diadromous fishes of U.K. conservation importance to be affected by MRED is reviewed. The information on which to base the review was found to be limited with respect to all aspects of these fishes' migratory behaviour and activity, especially with regards to MRED deployment, making it difficult to establish cause and effect relationships. The main findings, however, were that diadromous species can use the Earth's magnetic field for orientation and direction finding during migrations. Juveniles of anadromous brown trout (sea trout) Salmo trutta and close relatives of S. trutta respond to both the Earth's magnetic field and artificial magnetic fields. Current knowledge suggests that EMFs from subsea cables may interact with migrating Anguilla sp. (and possibly other diadromous fishes) if their movement routes take them over the cables, particularly in shallow water (

To investigate the ability of elasmobranchs to distinguish between differing prey-type electric fields we examined the electroreceptive foraging behaviour of a model species, Scyliorhinus canicula (small-spotted catshark). Catshark preferences were studied by behaviourally conditioning them to swim through narrow tunnels, and on exit presenting them simultaneously with two different electric fields. Their subsequent choices of the following paired options were recorded; (i) Two artificial electric fields (dipole electrodes) with different magnitude direct current (D.C.), (ii) Two artificial electric fields, one D.C. and the other alternating current (A.C.), of the same magnitude, and (iii) similar magnitude, natural and artificial D.C. electric fields associated with shore crabs and dipole electrodes respectively. We found a highly significant preference for the stronger D.C. electric field and a less pronounced, but still significant, preference for the A.C. electric field rather than the D.C. electric field. No preference was demonstrated between the artificial and natural D.C. electric fields. The findings are discussed in relation to the animal’s diet and ecology and with regard to anthropogenic sources of electric fields within their habitat.

Marine renewable energy promises to assist in the effort to reduce carbon emissions worldwide. As with any large-scale development in the marine environment, however, it comes with uncertainty about potential environmental impacts, most of which have not been adequately evaluated—in part because many of the devices have yet to be deployed and tested. We review the nature of environmental and, more specifically, ecological effects of the development of diverse types of marine renewable energy—covering marine wind, wave, tidal, ocean current, and thermal gradient—and discuss the current state of knowledge or uncertainty on how these effects may be manifested. Many of the projected effects are common with other types of development in the marine environment; for example, additional structures lead to concerns for entanglement, habitat change, and community change. Other effects are relatively unique to marine energy conversion, and specific to the type of energy being harnessed, the individual device type, or the reduction in energy in marine systems. While many potential impacts are unavoidable but measurable, we would argue it is possible (and necessary) to minimize others through careful device development and site selection; the scale of development, however, will lead to cumulative effects that we must understand to avoid environmental impacts. Renewable energy developers, regulators, scientists, engineers, and ocean stakeholders must work together to achieve the common dual objectives of clean renewable energy and a healthy marine environment.

To examine potential effects of male–female interactions on the sexes' foraging behaviour, we studied equal-sized, single- and mixed-sex groups of a benthic elasmobranch, the small-spotted catshark, in response to electric stimuli. A total of 88 size-matched, adult individuals were repeatedly presented with a range of artificial, prey type electric fields (E fields) under laboratory conditions in groups of (1) four males, (2) four females and (3) two of each sex. Females in single-sex groups were more responsive towards E fields than both single- and mixed-sex group males, despite showing similar activity levels. Equally, females in mixed-sex groups were more responsive than males in mixed groups, but showed similar activity levels. Furthermore, females in mixed-sex groups were less responsive to E fields than females from single-sex groups, despite showing similar activity levels. Males in mixed-sex groups were less responsive than males in single-sex groups, but were more active. These results indicate more intense foraging behaviour among females than males and significant reductions in foraging behaviour when grouped with the opposite sex. Both findings are probably associated with consequences of differing reproductive strategies and resultant sexual conflict (specifically male harassment and female avoidance).

Summary 1Global-scale environmental degradation and its links with non-renewable fossil fuels have led to an increasing interest in generating electricity from renewable energy resources. Much of this interest centres on offshore renewable energy developments (ORED). The large scale of proposed ORED will add to the existing human pressures on coastal ecosystems, therefore any ecological costs and benefits must be determined. 2The current pressures on coastal ecology set the context within which the potential impacts (both positive and negative) of offshore renewable energy generation are discussed. 3The number of published peer-review articles relating to renewable energy has increased dramatically since 1991. Significantly, only a small proportion of these articles relate to environmental impacts and none considers coastal ecology. 4Actual or potential environmental impact can occur during construction, operation and/or decommissioning of ORED. 5Construction and decommissioning are likely to cause significant physical disturbance to the local environment. There are both short- and long-term implications for the local biological communities. The significance of any effects is likely to depend on the natural disturbance regime and the stability and resilience of the communities. 6During day-to-day operation, underwater noise, emission of electromagnetic fields and collision or avoidance with the energy structures represent further potential impacts on coastal species, particularly large predators. The wider ecological implications of any direct and indirect effects are discussed. 7Synthesis and applications. This review demonstrates that offshore renewable energy developments will have direct and, potentially, indirect consequences for coastal ecology, with these effects occurring at different scales. Ecologists should be involved throughout all the phases of an ORED to ensure that appropriate assessments of the interaction of single and multiple developments with the coastal environment are undertaken.

Over a number of decades the process of prey choice has been investigated using fishes as model predators. Using fishes for the model has allowed the proximate factors that determine how a mobile predator finds and chooses to eat the prey encountered within a variable 3-D environment to be estimated. During prey choice a number of constraints exist, in particular most fish predators will eat their prey whole thus their jaws and gut create functional limitations once a prey has been attacked. By considering the relationship between the size of the prey and the predator's feeding apparatus and feeding motivation this study explores the link between mechanistic studies and theoretical, optimal foraging based predictions. How the prediction of prey choices made by the fish following prey encounter can be reconciled with what is likely to he found in the fish's stomach is discussed. This study uses a progression of empirical examples to illustrate how the limits of functional constraints and prey choice at different stages of motivation to feed can be taken into account to improve predictions of predator prey choice.

Summary 1 Climate is changing. Predictions are for at least a 3 °C rise in mean temperature in northern Europe over the next century. Existing severe impacts of nutrients and inappropriate fish stocking in freshwater systems remain. 2 Effects of warming by 3 °C above ambient, nutrient addition and the presence or absence of sticklebacks Gasterosteus aculeatus were studied in experimental microcosms dominated by submerged plants, mimicking shallow lake ecosystems. 3 Warming had considerably smaller effects on the phytoplankton community than did fish and nutrients. It had very minor effects on chlorophyll a and total phytoplankton biovolume. However, it significantly decreased the biovolumes of Cryptophyceae (a major component in the controls) and Dinophyceae. Contrary to expectation, warming did not increase the abundance of blue-green algae (cyanophytes). Warming decreased the abundances of Cryptomonas erosa (Cryptophyceae) and Oocystis pusilla (Chlorophycota) and increased those of two other green algae, Tetraedron minimum and Micractinium pusillum. It had no effect on a further 17 species that were predominant in a community of about 90 species. 4 Fish and nutrients, either together or separately, generally increased the crops of most of the 21 abundant species and of the algal groups. Exceptions were for diatoms and chrysophytes, which were very minor components of the communities. Fish, but neither nutrients nor warming, increased the number of species of phytoplankton detected. This was probably through removal of zooplankton grazers, and parallels terrestrial studies where the presence of top predators, by controlling herbivores, leads to increased plant diversity. 5 There was no particular pattern in the taxonomy or biological characteristics of those species affected by the treatments. In particular, there was no link between organism size (a surrogate for many important biological features of phytoplankton species) and the effects of warming, nutrient addition or presence or absence of fish. However, all species were relatively small and potentially vulnerable to grazing. 6 Synthesis and applications. The results suggest that fears of an increasing abundance of cyanophytes with current projections of global warming may be unrealized, at least in shallow unstratified lakes still dominated by macrophytes. However, they emphasize that eutrophication and fish manipulations remain very important impact factors that determine the abundance of phytoplankton and subsequent problems caused by large growths.

Top predators inhabiting a dynamic environment, such as coastal waters, should theoretically possess sufficient cognitive ability to allow successful foraging despite unpredictable sensory stimuli. The cognition-related hunting abilities of marine mammals have been widely demonstrated. Having been historically underestimated, teleost cognitive abilities have also now been significantly demonstrated. Conversely, the abilities of elasmobranchs have received little attention, despite many species possessing relatively large brains comparable to some mammals. The need to determine what, if any, cognitive ability these globally distributed, apex predators are endowed with has been highlighted recently by questions arising from environmental assessments, specifically whether they are able to learn to distinguish between anthropogenic electric fields and prey bioelectric fields. We therefore used electroreceptive foraging behaviour in a model species, Scyliorhinus canicula (small-spotted catshark), to determine cognitive ability by analysing whether elasmobranchs are able to learn to improve foraging efficiency and remember learned behavioural adaptations. Positive reinforcement, operant conditioning was used to study catshark foraging behaviour towards artificial, prey-type electric fields (Efields). Catsharks rewarded with food for responding to Efields throughout experimental weeks were compared with catsharks that were not rewarded for responding in order to assess behavioural adaptation via learning ability. Experiments were repeated after a 3-week interval with previously rewarded catsharks this time receiving no reward and vice versa to assess memory ability. Positive reinforcement markedly and rapidly altered catshark foraging behaviour. Rewarded catsharks exhibited significantly more interest in the electrical stimulus than unrewarded catsharks. Furthermore, they improved their foraging efficiency over time by learning to locate and bite the electrodes to gain food more quickly. In contrast, unrewarded catsharks showed some habituation, whereby their responses to the electrodes abated and eventually entirely ceased, though they generally showed no changes in most foraging parameters. Behavioural adaptations were not retained after the interval suggesting learned behaviour was not memorised beyond the interval. Sequences of individual catshark search paths clearly illustrated learning and habituation behavioural adaptation. This study demonstrated learning and habituation occurring after few foraging events and a memory window of between 12 h and 3 weeks. These cognitive abilities are discussed in relation to diet, habitat, ecology and anthropogenic Efield sources. © 2013 Springer-Verlag Berlin Heidelberg.

Following the construction of a large-scale public aquarium, we were presented with an opportunity to investigate how wild caught Caribbean reef fish respond to their first encounter with a novel environment. Within the constraints of this opportunity, we designed a behavioural study to determine the reef fishes' response to a new habitat in relation to their locomotory mode. Nine species of fish representing three locomotory modes: carangiform, sub-carangiform and labriform/sub-carangiform were observed over a four-week period following their first introduction to the aquarium. Fish activity levels and spatial distribution were quantified in relation to time since their first encounter with the novel environment. The most important result was that, regardless of locomotory mode or ecology, all of the species extensively explored the novel environment rather than settle on the first habitat that they encountered. This is a particularly interesting result for territorial species. More specifically, however, there were significant differences between species in activity through time. Carangiform activity level was lowest in the initial phases of an encounter with the novel environment subsequently rising to a stable level. The other species had variable activity throughout the study, but all of them exhibited a phase of low activity at some stage during the study. In terms of the fishes' use of the 2.5 million litres of water, six species utilised the whole of the aquarium based on a predefined zoning scheme. Although the initial activity level was low, carangiform swimmers used at least 90% of the zones in the early phases of an encounter with the novel environment and subsequently used all of the zones. Sub-carangiform species also used 100% of the zones by the end of the study. Three of the four labriform/sub-carangiform swimmers used a maximum of 90% of the zones. There was no significant difference between species in their use of the zones. However, each individual zone was subject to differential use by the fish. Owing to the extensive scale of the aquarium, we discuss the applicability of the behavioural results obtained to the natural environment in the context of the ecology of the species of fish studied.

When a three-spined stickleback Gasterosteus aculeatus encountered prey simultaneously the probability of hanging and the median pursuit time were greater than when prey were encountered sequentially. During simultaneous prey encounter fish did not choose to attack the more profitable prey but instead the nearer prey was handled first except when the difference between the two prey sizes was large. No difference was found in the level of total energetic intake by the fish regardless of prey size pairing. Fish that handled and ate the first prey of a pair in

A basic ecological tenet is that a predator chooses the prey that it consumes according tothe energetic gain available in relation to the time cost of dealing with each prey. For afish predator, the energetic content of prey increases with size whilst there is an associatedincrease in handling time (Werner, 1974; Kislalioglu & Gibson, 1976; Hoyle & Keast,1986; Gill & Hart, 1994). Prey choice can be based on the profitability of each size ofprey where the greatest profitability is defined as the prey size giving the maximum energygain per unit handling time (Stephens & Krebs, 1986). For a 45-mm

Changes in the foraging behaviour due to variation in the body size of the three-spined stickleback Gasterosteus aculeatus were investigated. All sizes of fish had a high probability of attacking prey whenever encountered. The probability of eating the prey increased with the size of the fish, as the larger fish had larger jaws and a greater stomach capacity. Therefore, as fish increased in size there was an increase in the probability of successful prey capture. The level of satiation did not have an effect on the prey handling time, which is contrary to other studies and is probably a result of the large prey sizes. The physical size of the prey meant that the handling times were long regardless of the motivational level of the fish. The larger fish took in more energy and at a faster rate, although the time to reach satiation was similar for all fish sizes. The advantage that large fish appear to have in successfully gaining large prey is negated by their greater metabolic requirement. The changes in feeding performance induced by small increases in body size could have important consequences for intraspecific competition, habitat Use and risk of predation.

Competition between pairs of stickleback was studied to investigate the effects of fish size and prey size on the feeding behaviour and competitive ability of the fish when single prey items were encountered sequentially. All the fish had a high probability of attacking the prey encountered. A fish with a high probability of being first to handle the prey had a high probability of ingesting it only when the prey was the smaller of the two prey sizes offered. With the larger prey, food gain was predicted not by the first to handle but by the jaw size of the fish. Encounter with a larger prey resulted in each fish attempting to capture it more often. This was due to the larger prey being visible for longer as it had to be manipulated before being swallowed. Even when prey were ingested and hence not visible, the unsuccessful competitor remained close by for approximately 7 s. The probability of eating any prey encountered was dependent on the phenotypic properties of the individual fish. A fast reacting fish with large jaws was able to reach its daily energy requirement regardless of the encounter rate with any prey size. Competitive ability was therefore a function of the fish, ability to reach the prey first, the capacity of the jaw apparatus to ingest the prey and prey encounter rate. When the pairs of fish encountered a high proportion of large prey, a generalist feeding strategy was the best option. The results can be applied to current foraging models, in particular to extending the sequential encounter model.

We describe 15 polymorphic dinucleotide microsatellite loci in the black-and-gold chromis, Neoglyphidodon nigroris (Cuvier: Teleostei: Pomacentridae). Microsatellites were isolated from a partial genomic library that was enriched for CA repeat motifs. The 15 loci yielded between two and 23 alleles per locus in a sample of 16 fish from two forereef sites off the island of Hoga, Wakatobi National Park, Indonesia. Observed and expected heterozygosities varied from 0.13 to 1.00 and 0.24 to 0.98, respectively. These markers should allow us to discriminate between closely related individuals and also to assess the connectedness of populations of N. nigroris that inhabit different reefs.

There is an urgent need to obtain information on the effects of underwater sound on marine fish due to imminent policy drivers, e.g., the European Union Marine Strategy Framework Directive, on one hand and the increasingly noisy activities in the marine environment on the other. Yet studying the influence of sound, particularly on the behavior of fish, is a challenging task. Studies in tanks can suffer problems with the reflection of sound, especially at the low frequencies that are most important for fish. Studies in the field are often limited because the observation of fish is very complicated.

Abstract. This study investigated changes in the feeding behaviour and prey choice of threespine stickleback, Gasterosteus aculeatus , of different body sizes, in response to increasing prey size and stomach fullness of the fish. Within the behavioural feeding sequence, a decision to attack and a decision to eat the prey were made by the fish. Smaller fish were more likely to stop in midwater during the attack and to manipulate the prey when handling it. Larger fish were more likely to attack and eat the prey. The probability of success, however, decreased with larger prey. Success also decreased with increasing stomach fullness when prey larger than the prey width:mouth width (PW:MW) ratio of 0·6 were encountered. Regardless of fish or prey size, there was a constant handling time of approximately 3 s during which the fish decided to eat. The time taken to handle and eat a prey decreased for larger fish and rose with larger prey. Spitting frequency increased with prey size and spitting was always required for prey greater than the 0·6 PW:MW ratio. This corresponded to a preference for prey oriented head first and ventral side up. Prey closest to the 0·6 PW:MW ratio were the largest prey that could be eaten with little change in the time cost over all levels of stomach fullness. These prey, therefore, gave the fish the best energy return per unit cost, except when the stomach was empty, as larger prey represented a bigger energy return for the same time cost giving them a greater effective profitability. The morphological relationship between a predator and prey determines the resulting feeding behaviour and prey choice, although this choice also depends on the predator's need to acquire food.

(1993). Choosing prey size: A comparison of static and dynamic foraging models for predicting prey choice by fish. Marine Behaviour and Physiology: Vol. 23, Behavioural Ecology of Fishes, pp. 91-104.

An experiment was designed to study how gut fullness and encounter with 5-mm Asellus aquaticus influenced acceptance or rejection of less profitable 8-mm Asellus. 45-mm sticklebacks were found to always accept 5-mm prey whereas 8-mm prey were accepted with an initial probability of about 0.9. This probability decreased as the gut filled. Fish of differing sizes and sex had similar daily energy intakes per unit body size, however the acceptance of 8-mm prey was related to fish size. Whenever a fish orientated to a prey it was followed by pursuit and manipulation independently of prey size. The decision to accept or reject prey occurred after one manipulation, a criterion that was more variable for the larger prey. For one feeding session per day the total energy intake was almost constant despite the changing combination of prey sizes eaten. The fish ate prey with long handling times if the energetic contents of the stomach had not reached 450 J. Calculations were made of how many of each millimetre prey size group would satisfy the 450 J demand and how long the estimated number would take to handle. This showed that the best option is to consume 5-mm prey if given the choice.

Since the mass mortality of the urchin Diadema antillarum in 1983, parrotfishes have become the dominant grazer on Caribbean reefs. The grazing capacity of these fishes could be impaired if marine reserves achieve their long-term goal of restoring large consumers, several of which prey on parrotfishes. Here we compare the negative impacts of enhanced predation with the positive impacts of reduced fishing mortality on parrotfishes inside reserves. Because large-bodied parrotfishes escape the risk of predation from a large piscivore (the Nassau grouper), the predation effect reduced grazing by only 4 to 8%. This impact was overwhelmed by the increase in density of large parrotfishes, resulting in a net doubling of grazing. Increased grazing caused a fourfold reduction in the cover of macroalgae, which, because they are the principal competitors of corals, highlights the potential importance of reserves for coral reef resilience.