Failure of predator conditioning: an experimental study of predator avoidance in brown trout (Salmo trutta)

Many studies have documented that hatchery‐reared salmonids generally have inferior survival after being stocked compared with wild conspecifics, hatchery and wild salmonids have been observed to differ in their antipredator responses. The response of brown trout (Salmo trutta) juveniles (0+) of differing backgrounds to a live predator was compared in two experiments. First, the antipredator behaviour of predator‐naïve hatchery‐reared brown trout and wild‐exposed brown trout were assessed in behavioural trials which lasted for eight days. Second, predator‐naïve and predator‐conditioned hatchery‐reared brown trout were assessed in identical behavioural trials. Brown trout were ‘predator‐conditioned’ by being held in a stream‐water aquarium with adult Atlantic salmon (Salmo salar) and adult brown trout for two days prior to behavioural trials. Predator‐conditioned hatchery‐reared brown trout spent more time in shelters in the trial aquaria than predator‐naïve hatchery‐reared fish, but did not differ in time spent in the predator‐free area. Predator conditioning may account for the increased time spent in the shelter, but does not appear to have affected time spent in the predator‐free area. However, even if significant alteration in behaviour can be noted in the laboratory, the response might not be appropriate in the wild.     

Interactions between South American sea lions Otaria flavescens (Shaw) and salmon farms in southern Chile

By means of surveys in 48 salmon farms in Chile, and the analysis of fish mortality in 23, the results showed that attacks by the South American sea lion, Otaria flavescens, were periodical at a daily and seasonal scale, and because of their natural nocturnal feeding behaviour, were more frequent at night and in fall–winter seasons. The attacks varied markedly between the farms studied. No relation was found between the intensity of attacks, the distance to the nearest colony of sea lions and the population in the colony. Differences found between farms could be because of the different protective devices used, namely anti‐predator nets, acoustic harassment devices (AHDs) and fibreglass models of killer whales, and the way in which they were used. Of these devices, only the anti‐predator net achieved a permanent reduction in sea lion attacks. Both AHDs and fibreglass killer whales were inefficient in the short term.     

Effects of simulated light intensity,habitat complexity and forage type on predator–prey interactions in walleye Sander vitreus

Predator‐prey interactions can be influenced by the behaviour of individual species as well as environmental factors. We conducted laboratory experiments to test for the influences of two abiotic factors (light intensity and habitat complexity) on predator–prey interactions between walleye Sander vitreus and two prey species, bluegill Lepomis macrochirus and golden shiner Notemigonus crysoleucas. Three light intensities were simulated (day, twilight and night) in the presence or absence of simulated vegetation. Observations of predator behaviour indicated that walleye increased activity and foraging success with decreasing light levels and had most success capturing dispersed, closer prey. While schooling could not be maintained as light levels diminished, prey decreased predation vulnerability by moving into vegetation or higher in the water column. Throughout all treatments, bluegill were more evasive to capture as the number of strikes was similar on both prey but capture rates were higher for golden shiner. Although light intensity and simulated habitat complexity affected predator and prey behaviour, these factors did not interact to influence foraging success of walleye. To fully understand predator and prey behaviours in fishes, an understanding of species‐specific responses to abiotic and biotic factors is necessary.     

Are hatchery‐reared abalone naïve of predators? Comparing the behaviours of wild and hatchery‐reared northern abalone,Haliotis kamtschatkana (Jonas, 1845)

Abalone populations have declined worldwide, generating interest in enhancement using hatchery‐reared individuals. In many cases, such restoration efforts have met with limited success due to high predator‐induced mortality rates. Furthermore, the mortality rates of outplanted hatchery abalone are often considerably higher than for wild individuals. This study uses northern abalone (Haliotis kamtschatkana) as a case study to determine whether hatchery‐reared abalone behave differently than their wild counterparts. In the field, outplanted hatchery‐reared abalone were significantly less responsive than wild abalone, in terms of number of abalone responding and intensity of response, to nearby movement and to physical contact with an inert probe. Also, when encountering a cue to which all abalone responded (a seastar predator), hatchery‐reared individuals remained subdued. Anti‐predator behavioural deficits in hatchery‐reared abalone were more pronounced in 4‐year‐old individuals than in 1‐year‐old individuals, suggesting an influence of either age or amount of time spent in the hatchery environment. These behavioural differences are expected to increase the vulnerability of hatchery‐reared abalone to predators, and are likely a major cause of their elevated predator‐induced mortality when outplanted.     

Foraging behaviour of juvenile pink salmon (Oncorhynchus gorbuscha) and size‐dependent predation risk

Two hypotheses related to effects of juvenile pink salmon (Oncorhynchus gorbuscha) foraging behaviour and size on their predation risk were evaluated using field data collected in Prince William Sound, Alaska 1 995–97. My results supported the hypothesis that low macrozooplankton density leads to dispersion of juvenile salmon from shallow nearshore habitats and greater predation risk, but zooplankton type was an important factor. When the biomass of large copepods (primarily Neocalanus spp.) declined, salmon dispersed from shallow nearshore habitats, and mean daily individual predator consumption of salmon increased by a factor of 5. A concomitant five‐fold increase in the probability of occurrence of salmon in predator stomachs supported the notion that increased predation on salmon was caused by a greater overlap between predator and prey when salmon dispersed offshore, not an increase in the number of salmon consumed per feeding bout. The results also generally supported the hypothesis that the timing of predation events modifies the nature of size‐dependent predation losses of salmon to different predator groups (small and large planktivores and piscivores). Size‐dependent vulnerabilities of salmon to predators were a function of both predator and prey sizes. When simulated predation was shifted from May to June, the vulnerability of salmon became more dependent on their growth than initial size. But, the size‐ and growth‐dependent vulnerabilities of salmon differed more among predator groups than between May and June, suggesting that changes in the composition of predator fields could more strongly affect the nature of size‐dependent predation than changes in the timing of predation losses.     

Do intracoelomic telemetry transmitters alter the post‐release behaviour of migratory fish?

Electronic tags have become a common tool in fish research, enhancing our understanding of how fish interact with their environment and move among different habitats, for estimating mortality and recording internal physiological states. An often‐untested assumption of electronic tagging studies is that tagged fish are representative of untagged conspecifics and thus show ‘normal’ behaviour (e.g. movement rates, swimming activity, feeding). Here, we use a unique data set for potamadromous walleye (Sander vitreus) in Lake Huron and Lake Erie tributaries to assess whether the lack of appropriate controls in electronic tagging could seriously affect behavioural data. We used fish tagged in previous years and compared their migratory behaviour during the spawning season to fish tagged in a current year at the same location. The objective of the study was to determine whether intracoelomic acoustic tag implantation altered downstream movement of walleye after spawning. Fish tagged in a given season travelled slower downstream from two river spawning sites than fish tagged in previous years. Fish tagged one or two years earlier showed no differences between each other in downstream travel time, in contrast to fish tagged in a given year. Our results support notions that standard collection and intracoelomic tagging procedures can alter short‐term behaviour (i.e. days, weeks, months), and as such, researchers should use caution when interpreting data collected over such time periods. Further, whenever possible, researchers should also explicitly evaluate post‐tagging effects on behaviour as part of their experimental objectives.     

Impacts of domestication on angling vulnerability of common carp,Cyprinus carpio: the role of learning,foraging behaviour and food preferences

Domestication in fish selection increases vulnerability to angling. Two common garden‐reared genotypes of common carp, Cyprinus carpio L., differing in degree of domestication (highly domesticated mirror carp and less domesticated scaled carp) were exposed to fishing in two environments (i.e. ponds and laboratory tanks) to quantify vulnerability to angling. Foraging behaviour and food preferences were quantified to explain variation in angling vulnerability in a mechanistic manner. Domesticated mirror carp were more vulnerable to angling gear than scaled carp in both environments, which was related to greater food intake and bolder‐foraging behaviour. Independent of genotype, catchability decreased and time until first capture increased over fishing time, indicating learned hook avoidance. No differences were observed in food preferences among genotypes, rendering bait‐selective feeding an unlikely explanation for differential vulnerability to angling. It was concluded that vulnerability to angling has a genetic basis in carp and that boldness plays a paramount role in explaining why more domesticated genotypes are more easily captured by angling.     

Size‐dependent stress response in juvenile Arctic charr (Salvelinus alpinus) under prolonged predator conditioning

Predator conditioning can be used to improve post‐release antipredator recognition of hatchery‐reared salmonids. However, possible negative stress‐related effects of prolonged predator conditioning on juvenile fish physiology are poorly understood. We studied the effects of prolonged (91 days) predator odour exposure on whole‐body cortisol level and spleen size in six full‐sib families of juvenile hatchery‐bred Arctic charr (Salvelinus alpinus). Chemical cues from water containing charr‐fed pikeperch (Sander lucioperca) were used as the predator exposure stimuli and lakewater was used as a chemical control. Our study revealed that juvenile body cortisol levels post‐predator conditioning were affected by treatment, fish size and their interaction. Importantly, among the smaller (i.e. slowest growing) charr, the predator‐exposed fish had higher cortisol levels than control fish, while the opposite pattern was true for the larger fish. These results suggest that chemical cues from charr‐fed predators induce a prolonged stress response in juvenile charr. As prolonged predation exposure seems to elevate stress levels in a size‐dependent manner, the larger, faster growing fish could possibly have intrinsically lower stress responses to predation threats than smaller, slower growing fish. Possible coupling between stress sensitivity and growth requires further attention due to the likely implications for the management of unintended domestication among captive‐reared salmonids.     

Rapid dispersion of escaped meagre (Argyrosomus regius) from a coastal Mediterranean fish farm

Meagre (Argyrosomus regius) is an emerging species in aquaculture considered a locally absent species in the Western Mediterranean. Little is known about meagre escape incidents from farms. To evaluate escape impacts and develop mitigation measures, knowledge about the behaviour of escaped meagre is necessary. In this study, a rapid dispersion of escaped meagre was observed following a simulated escape incident in a coastal Mediterranean farm, using acoustic telemetry (acoustic tag: AT) and mark‐and‐recapture techniques (external tag: ET). A small proportion of AT meagre (22.7%) remained within the farm boundaries 24 h after the simulated escape. A total of 84 meagre (9.2%) were captured by local trammel netters nearby the farm facility within the first 48 h after escape. Only two AT individuals (15.3%) were detected by acoustic receivers located in coastal areas within 2 days after dispersal from the farm and eight ET meagre (0.8%) were recaptured along the shoreline during the first week. Two AT individuals (15.3%) were considered to be dead by predation few hours after escape. The rest of the AT individuals (N = 6, 46.2%) left the facility alive, and they were never detected by receivers nor reported back by fishermen. Thus, our results highlight the necessity of establishing escape mitigation strategies at Mediterranean fish farms, i.e. recapture programmes 24–48 h after the escape incident in collaboration with local fishermen, to diminish potential impacts related to meagre escapees.     

Learning about danger: chemical alarm cues and local risk assessment in prey fishes

An individual's behaviour patterns can be conceptualized as a series of threat‐sensitive trade‐offs between ambient predation pressure and a suite of fitness‐related activities, such as resource defence, foraging and mating. Individuals that can reliably assess local predation risk could increase their fitness potential by exhibiting predator avoidance behaviours only at appropriate times. However, such learned risk assessment requires reliable information regarding current predation risks. A diverse range of prey fishes are known to possess chemical alarm cues, which when detected by conspecifics and some heterospecifics, elicit a variety of overt and covert responses. These chemical cues, either alone or as a part of a predator's dietary odour, provide reliable information regarding local predation risk. In this review, I describe recent works examining the role of chemosensory information in: (i) acquired predator recognition, (ii) predator inspection behaviour and (iii) the use of conspecific and heterospecific cues as social information sources.     

Intercohort cannibalism and parturition-associated behaviour of adult swordtail, Xiphophorus helleri Heckel (Pisces: Poeciliidae)

The post‐parturition behavioural patterns of adult Xiphophorus helleri (Heckel) were described and quantified by recording their duration and/or frequency in glass tanks. All births occurred in the dark. Female behaviour might increase the chance of newborn juvenile survival: at the top of the tank, where juveniles are more vulnerable to predation, the females gave birth in the artificial refuge, while no parturition took place in the refuge at the bottom. Birth‐giving females were no less cannibalistic than males and other females. There was a reduction in cannibalism and the frequency of secondary attacks with time, even though the number of juveniles in the tank continued to increase due to ongoing parturition. Most adults were found at the bottom of the water column during the first 500 min of parturition, after which they were distributed equally in the water column. The movement of adults, away from the bottom and consequently away from the juveniles corresponded with an increase in feeding on alternative food and a reduction in cannibalism. Thus, cannibalism in swordtail might be opportunistic predation.     

Variation in wind and piscivorous predator fields affecting the survival of Atlantic salmon,Salmo salar,in the Gulf of Maine

Abstract Observations relevant to the North American stock complex of Atlantic salmon, Salmo salar L., suggest that marine mortality is influenced by variation in predation pressure affecting post‐smolts during the first months at sea. This hypothesis was tested for Gulf of Maine (GOM) stocks by examining wind pseudostress and the distribution of piscivorous predator fields potentially affecting post‐smolts. Marine survival has declined over recent decades with a change in the direction of spring winds, which is likely extending the migration of post‐smolts by favouring routes using the western GOM. In addition to changes in spring wind patterns, higher spring sea surface temperatures have been associated with shifting distributions of a range of fish species. The abundance of several pelagic piscivores, which based on their feeding habits may predate on salmon post‐smolts, has increased in the areas that serve as migration corridors for post‐smolts. In particular, populations of silver hake, Merluccius bilinearis (Mitchell), red hake, Urophycis chuss (Walbaum), and spiny dogfish, Squalus acanthias L., increased in size in the portion of the GOM used by post‐smolts. Climate variation and shifting predator distributions in the GOM are consistent with the predator hypothesis of recruitment control suggested for the stock complex.     

Escape‐related behaviour of Atlantic cod,Gadus morhua L., in a simulated farm situation

Cultured fish escaping from farms represent an economic loss as well as a potential problem for wild fish populations. This study investigated the escape‐related behaviour of farmed Norwegian coastal and northeast Arctic cod (NEAC), Gadus morhua L. Six groups of fish were observed during three replicate trials in a large tank that was split into two equal parts by a net wall. The fish could move freely through an opening in the net during the trials. Three groups were not fed during the trials. The first escape occurred shortly after producing the net opening (11±10.5 min; mean±standard deviation). Norwegian coastal cod were more prone to escape than the NEAC. A starvation period of 9 days increased the number of fish on the escape side of the tank. Net biting and net inspection by the fish were frequently observed, irrespective of whether the fish were fed or not. The same fish were repeatedly found on the escape side of the tank, but the propensity for recurrent escape behaviour was not related to genotype, feeding status or size. The results suggest that other factors, such as individual variation in boldness or exploration behaviour, could affect the willingness to escape.     

Spatio‐temporal analyses of marine predator diets from data‐rich and data‐limited systems

Accounting for variation in prey mortality and predator metabolic potential arising from spatial variation in consumption is an important task in ecology and resource management. However, there is no statistical method for processing stomach content data that accounts for fine‐scale spatio‐temporal structure while expanding individual stomach samples to population‐level estimates of predation. Therefore, we developed an approach that fits a spatio‐temporal model to both prey‐biomass‐per‐predator‐biomass data (i.e. the ratio of prey biomass in stomachs to predator weight) and predator biomass survey data, to predict “predator‐expanded‐stomach‐contents” (PESCs). PESC estimates can be used to visualize either the annual landscape of PESCs (spatio‐temporal variation), or can be aggregated across space to calculate annual variation in diet proportions (variation among prey items and among years). We demonstrated our approach in two contrasting scenarios: a data‐rich situation involving eastern Bering Sea (EBS) large‐size walleye pollock (Gadus chalcogrammus, Gadidae) for 1992–2015; and a data‐limited situation involving West Florida Shelf red grouper (Epinephelus morio, Epinephelidae) for 2011–2015. Large walleye pollock PESC was predicted to be higher in very warm years on the Middle Shelf of the EBS, where food is abundant. Red grouper PESC was variable in north‐western Florida waters, presumably due to spatio‐temporal variation in harmful algal bloom severity. Our approach can be employed to parameterize or validate diverse ecosystem models, and can serve to address many fundamental ecological questions, such as providing an improved understanding of how climate‐driven changes in spatial overlap between predator and prey distributions might influence predation pressure.     

Cumulative selectivity benefits of increasing mesh size and using escape gaps in Australian Portunus armatus traps

The individual and cumulative effects of increasing mesh size (from 56‐ to 75‐mm stretched mesh opening) and installing three escape gaps (36 × 120 mm) in collapsible‐netted round traps were assessed to address concerns associated with excessive discarding in an Australian portunid fishery. Compared to conventional traps comprising 56‐mm mesh throughout, those with the same mesh size and escape gaps caught significantly fewer (by 54%) undersized blue swimmer crabs, Portunus armatus Milne‐Edwards and yellowfin bream, Acanthopagrus australis Günther (by 64%). Irrespective of escape gaps, simply increasing the mesh size to 75 mm did not significantly affect catches of undersized P. armatus, although 87% fewer A. australis were retained. Traps with both 75‐mm mesh and escape gaps maintained reductions of A. australis, but had a clear cumulative effect on P. armatus selection, retaining 84% fewer undersized individuals across a larger size at retention. The results support using escape gaps in existing conventional traps, but illustrate the need to configure the minimum legal mesh size to approach the desired target size of P. armatus as a precursor to maximising trap selectivity. Larger‐meshed traps also require less material (i.e. plastic), which benefits their manufacture and, equally importantly, reduces environmental costs when lost.     

Multispecies functional responses reveal reduced predation at high prey densities and varied responses among and within trophic groups

Understanding predator–prey interactions is critical for marine fisheries and ecosystem management as they shape community structure, regulate prey populations and present energy demands critical for community sustainability. We examined multispecies functional responses of 17 fishes (48 predator‐size combinations) spanning piscivores, planktivores and benthivores for the northeast US continental shelf. Similar to previous work, linear relationships between predation and prey density (Holling type I response) were not supported, since model estimates of handling time were greater than zero for the prey considered. Instead, a clear majority of the predators sampled were Holling type III feeders (sigmoidal; prey switching or learning). For piscivores, nearly all responses were Holling type III with the exception of one being Holling type II (hyperbolic; satiation). Planktivores and benthivores exhibited a combination of type IV (feeding confusion at high prey density) and Holling type III responses. The relationships were predator‐ and prey‐dependent, which is counter to assumptions that are often made of trophic groups. Decreased predation at high prey densities (type IV response) present among planktivores and benthivores suggests an overestimation of predation can occur if ignored. This contrasts with fish and squid prey which primarily invoked a Holling type III response. Functional responses are key to modelling trophic interactions for multispecies and ecosystem models. By refining these inputs in a multispecies context with empirical data, we can advance our understanding of whole‐shelf ecology and improve decision‐making tools for resource management.     

Vertical movements of bigeye tuna (Thunnus obesus) associated with islands, buoys, and seamounts near the main Hawaiian Islands from archival tagging data

To learn more about the movement patterns of bigeye tuna (Thunnus obesus), we deployed archival tags on 87 fish ranging in fork length from 50 to 154 cm. Thirteen fish were recaptured, from which 11 archival tags were returned, representing in aggregate 943 days‐at‐liberty. We successfully retrieved data from 10 tags, representing 474 days in aggregate. The largest fish recaptured was 44.5 kg [131 cm fork length (FL)] and the smallest 2.8 kg (52 cm). The deepest descent recorded was 817 m, the coldest temperature visited 4.7°C, and minimum oxygen level reached ～1 mL L^?1. Fish spent little time at depths where water temperatures were below 7°C and oxygen levels less than ～2 mL L^?1. Five fish were recaptured near the offshore weather buoy where they were tagged. Based on vertical movement patterns, it appeared that all stayed immediately associated with the buoy for up to 34 days. During this time they remained primarily in the uniform temperature surface layer (i.e. above 100 m). In contrast, fish not associated with a floating object showed the W‐shaped vertical movement patterns during the day characteristic of bigeye tuna (i.e. descending to ～300–500 m and then returning regularly to the surface layer). Four fish were tagged and subsequently recaptured near Cross Seamount up to 76 days later. These fish exhibited vertical movement patterns similar to, but less regular than, those of fish not associated with any structure. Bigeye tuna appear to follow the diel vertical movements of the deep sound scattering layer (SSL) organisms and thus to exploit them effectively as a prey resource. Average night‐time depth was correlated with lunar illumination, a behaviour which mimics movements of the SSL.     

Environmental noise in spawning areas: the case of Atlantic Bluefin Tuna (Thunnus thynnus)

This paper provides an exploratory analysis aiming to seek whether the colour of environmental noise theory could help in understanding the intriguing reproductive strategy of Atlantic bluefin tuna (BFT). A frequency‐based approach based on spectral exponents, f^β with β < 0, is chosen and applied on 10 biogeographical provinces covering the North Atlantic. The major BFT spawning area, i.e. the Mediterranean Sea, was the only one to display a pink power spectrum, whereas open ocean regions displayed more reddened fluctuations, i.e. greater variance at low frequencies. Environmental noise in the Mediterranean could, thus, offer more favourable characteristics on the long‐term than the open ocean. The implications of these findings are discussed in regards to medium and long (possibly evolutionary) time scales.     

Modelling transport of chokka squid (Loligo reynaudii) paralarvae off South Africa: reviewing,testing and extending the ‘Westward Transport Hypothesis’

Annual landings of chokka squid (Loligo reynaudii), an important fishing resource for South Africa, fluctuate greatly, and are believed to be related to recruitment success. The ‘Westward Transport Hypothesis’ (WTH) attributes recruitment strength to variability in transport of newly hatched paralarvae from spawning grounds to the ‘cold ridge’ nursery region some 100–200 km to the west, where oceanographic conditions sustain high productivity. We used an individual‐based model (IBM) coupled with a 3‐D hydrodynamic model (ROMS) to test the WTH and assessed four factors that might influence successful transport – Release Area, Month, Specific Gravity (body density) and Diel Vertical Migration (DVM) – in numerical experiments that estimated successful transport of squid paralarvae to the cold ridge. A multifactor ANOVA was used to identify the primary determinants of transport success in the various experimental simulations. Among these, release area was found to be the most important, implying that adult spawning behaviour (i.e., birth site fidelity) may be more important than paralarval behaviour in determining paralarval transport variability. However, specific gravity and DVM were found to play a role by retaining paralarvae on the shelf and optimizing early transport, respectively. Upwelling events seem to facilitate transport by moving paralarvae higher in the water column and thus exposing them to faster surface currents.     

Migration pathways,speed and mortality of Atlantic salmon (Salmo salar) smolts in a Scottish river and the near‐shore coastal marine environment

Long‐distance migration of Atlantic salmon (Salmo salar) is known to result in high levels of mortality. For a species experiencing global population decline, it is thus vital to better understand migration behaviour, both in the river and marine stages. Atlantic salmon smolts (n = 50) were tracked using acoustic telemetry in the River Deveron, Scotland, and adjacent coastal area. Higher rates of mortality were observed in the river (0.77% per km) than the early marine stage of migration (0.0% per km). Mortality likely resulted from predation. Higher swim speeds were recorded in the early marine stage compared with the river (marine = 7.37 ± 28.20 km/day; river = 5.03 ± 1.73 km/day [mean ± SD]), a potential predator avoidance behaviour. The majority of smolts leaving the river did so in darkness and on a flooding tide. Overall river and marine migration success were linked to nights of lower lunar brightness. Marine migration speed decreased with increasing environmental noise levels, a finding with implications for fisheries management. The migration pathway in the early marine environment did not follow obvious geographical features, such as the coastline. Thus, we suggest that early marine environment pathways are more influenced by complex water currents. These findings highlight factors that influence smolt migration survival and behaviour, areas on which future research should focus.