Foraging arena theory

There is a critical need for quantitative models that can help evaluate trade‐off decisions related to the impacts of harvesting and protection of aquatic ecosystems within an ecosystem context. Ecosystem models used to evaluate such trade‐offs need to have the capability of capturing the dynamic stability that can arise when predator‐prey interactions are restricted to spatial and temporal arenas. Foraging arenas appear common in aquatic systems and are created by a wide range of mechanisms, ranging from restrictions of predator distributions in response to predation risk caused by their own predators, to risk‐sensitive foraging behaviour by their prey. Foraging arenas partition the prey in each predator‐prey interaction in a food web into vulnerable and invulnerable states, with exchange between these states potentially limiting overall trophic flow. Inclusion of vulnerability exchange processes in models for recruitment processes and food web responses to disturbances like harvesting leads to very different predictions about dynamic stability, trophic cascades and maintenance of ecological diversity than do models based on large‐scale mass action (random mixing) interactions between prey and predators. Although a number of methods to estimate these critical exchange rates are presented, none are considered fully satisfactory. The most important challenge for the practical application of models that incorporate foraging arena theory today is not only developing new or improved methods for measuring exchange rates but also evaluating how such rates vary in responses to major fishery‐induced changes in abundances of predators.     

Trophic role of Atlantic cod in the ecosystem

As the world's oceans continue to undergo drastic changes, understanding the role of key species therein will become increasingly important. To explore the role of Atlantic cod ( Gadus morhua Gadidae) in the ecosystem, we reviewed biological interactions between cod and its prey, predators and competitors within six ecosystems taken from a broad geographic range: three are cod-capelin ( Mallotus villosus Osmeridae) systems towards cod's northern Atlantic limit (Barents Sea, Iceland and Newfoundland–Labrador), two are more diverse systems towards the southern end of the range (North Sea and Georges Bank–Gulf of Maine), and one is a species-poor system with an unusual physical and biotic environment (Baltic Sea). We attempt a synthesis of the role of cod in these six ecosystems and speculate on how it might change in response to a variety of influences, particularly climate change, in a fashion that may apply to a wide range of species. We find cod prey, predators and competitors functionally similar in all six ecosystems. Conversely, we estimate different magnitudes for the role of cod in an ecosystem, with consequently different effects on cod, their prey and predator populations. Fishing has generally diminished the ecological role of cod. What remains unclear is how additional climate variability will alter cod stocks, and thus its role in the ecosystem.     

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.     

Size-dependent predation on marine fish larvae by Ctenophores, Scyphomedusae, and Planktivorous fish

Size-dependent predation rates on marine fish larvae by the ctenophore Mnemiopsis leidyi , scyphomedusa Chrysaora quinquecirrha , and planktivorous fish Anchoa mitchilli were estimated via experiments in 3.2 m³ me-socosms. Predation rates on larvae of the goby Gobio-soma bosci were estimated in relation to 1) length of larval prey, 2) presence or absence of alternative < 1 mm zooplankton prey, and 3) a predator-prey interaction between the scyphomedusa and ctenophore. The consumption rate of larvae by the three predators generally declined as larval length increased. The ctenophore ate fewer (1.0 d^-1 per predator) larvae than did the medusa (7.8 d ^-1 per predator) or bay anchovy (4.0 d ^-1 per predator), but it consumed larvae in all size classes tested (3.0–9.5 mm SL). Predation by bay anchovy and the medusa was more size-dependent: 3.0–5.0 mm larvae were most vulnerable. Fewer larvae were eaten by the ctenophore and bay anchovy when zooplankton alternative prey was available, but predation on larvae by the medusa was not influenced by alternative zooplankton prey. Consumption rate of fish larvae by the medusa was reduced 20–25% when the ctenophore was present as its alternative prey. An encounter-rate model was developed and its parameters estimated from the experimental results. Model simulations indicated that the relationship between larval size and vulnerability is dependent on size, swimming speed, and behavior of both predators and larvae, and that bigger or faster-growing fish larvae are not always less vulnerable to predation.     

Ecological and Ethological Perspectives in Larval Fish Feeding

ABSTRACT

Low survival rates during larval stages constitute a major bottleneck in the successful culture of many marine and some freshwater fish. The availability of live food is recognized as a critical factor influencing larval survival. Live food is still superior to the best larval diets in terms of larval survival and growth. This paper reviews important ecological and ethological aspects of feeding, from hatching tothe weaning stage, and relates them to problems in larval culture. In general, freshwater fish larvae are easier to raise than marine fish larvae, because at hatching they are larger and endowed with more yolk reserves, are less sensitive to starvation, and canbe weaned to artificial diets sooner. The feeding behavior of the larvae can be analyzed in terms of the sequential components of predation: search, encounter, pursuit, attack, capture, and ingestion. The searching efficiency and encounter rates of the visual predator are influenced by prey parameters such as body size, conspicuousness, and evasiveness. Turbidity of the water and light intensity also affect prey detection. To changing prey densities, the larvae show typical Type II functional responses, which are influenced by prey handling time, which in turn is largely a function of prey size. Knowledge of larval functional responses is helpful in providing the right concentrations oflive food for larval culture. The larvae are initially gape-limited and exhibit prey size selectivity but gradually widen their prey size range as they grow. An aquacultural application of this is the commonly employed feeding protocol, prey size sequencing, in which progressively larger live food items are offered as the larvae grow. A thorough knowledge of the feeding behavior is also essential in the formulation of acceptable larval diets.     

Effects of predator species,vegetation and prey assemblage on prey preferences of predators with emphasis on vulnerability of age‐0 common carp

Abstract Predation is an important force structuring aquatic communities, but predator–prey interactions are complex and regulated by multiple factors. Invasive fishes may interact with native fishes to alter predator–prey preferences and community dynamics. For example, common carp, Cyprinus carpio L., is an invasive species that can become abundant and negatively affect aquatic ecosystems. Juvenile common carp are occasionally found in predator diets, but predator preferences for common carp compared with alternative prey remains unknown. Prey selection and feeding behaviour of five piscivores (flathead catfish, Pylodictis olivaris (Rafinesque); largemouth bass, Micropterus salmoides (Lacepède); smallmouth bass, M. dolomieu Lacepède; walleye, Sander vitreus (Mitchill); and northern pike, Esox lucius L.) foraging on juvenile common carp and two alternative prey (fathead minnow, Pimephales promelas Rafinesque, and yellow perch, Perca flavescens Mitchill) at variable densities and habitats were evaluated. Common carp and fathead minnow were generally selected for or neutrally selected across predator species, habitat types and prey assemblages. By contrast, yellow perch was generally selected against. Common carp were easily captured but difficult to manipulate and ingest compared with other prey. These results reveal that common carp are vulnerable to a variety of predators, suggesting control of this detrimental invader may be possible through biomanipulation.     

No room for dessert: A mechanistic model of prey selection in gut‐limited predatory fish

Predatory fish structure communities through prey pursuit and consumption and, in many marine systems, the gadoids are particularly important. These predators have flexible feeding behaviours and often feed on large prey items. Digestion times of large prey are usually longer than handling times, and gut processing limits feeding rate at high prey density. Optimizing the gut content mix can therefore be an important behavioural strategy. Here, we develop a foraging model that incorporates gut processing and use the model to disentangle internal and external limitations on feeding in the omnivorous cod (Gadus morhua, Gadidae). We extend the traditional definition of prey profitability to consider prey digestive quality, which we quantify for prey of Northeast Atlantic cod populations. We find an important role for gut limitation; within a range of ecologically relevant temperatures and prey densities, predicted feeding rates were strongly reduced compared to feeding constrained by external factors only, and the optimal diet composition under gut limitation differed from predictions from traditional foraging theory. Capelin, a main prey of Northeast Arctic cod, had the highest digestive quality of all prey across ecosystems, but the cold temperatures in the Barents Sea strongly limited feeding rate by slowing down digestion. Baltic cod fed on a higher proportion of poor‐quality prey compared to the other populations, contributing to its slow growth in relation to water temperature. Gut limitation is particularly important to consider in foraging models for fish with many alternative prey species or fish occupying cold waters where digestion is slow.     

Multiple predator effects and native prey responses to two non‐native Everglades cichlids

Abstract – Non‐native predators may have negative impacts on native communities, and these effects may be dependent on interactions among multiple non‐native predators. Sequential invasions by predators can enhance risk for native prey. Prey have a limited ability to respond to multiple threats since appropriate responses may conflict, and interactions with recent invaders may be novel. We examined predator–prey interactions among two non‐native predators, a recent invader, the African jewelfish, and the longer‐established Mayan cichlid, and a native Florida Everglades prey assemblage. Using field enclosures and laboratory aquaria, we compared predatory effects and antipredator responses across five prey taxa. Total predation rates were higher for Mayan cichlids, which also targeted more prey types. The cichlid invaders had similar microhabitat use, but varied in foraging styles, with African jewelfish being more active. The three prey species that experienced predation were those that overlapped in habitat use with predators. Flagfish were consumed by both predators, while riverine grass shrimp and bluefin killifish were eaten only by Mayan cichlids. In mixed predator treatments, we saw no evidence of emergent effects, since interactions between the two cichlid predators were low. Prey responded to predator threats by altering activity but not vertical distribution. Results suggest that prey vulnerability is affected by activity and habitat domain overlap with predators and may be lower to newly invading predators, perhaps due to novelty in the interaction.     

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.     

Turbidity‐induced changes in emergent effects of multiple predators with different foraging strategies

Abstract – In natural systems, prey frequently interact with multiple predators and the outcome often cannot be predicted by summing the effects of individual predator species. Multiple predator interactions can create emergent effects for prey, but how those change across environmental gradients is poorly understood. Turbidity is an environmental factor in aquatic systems that may influence multiple predator effects on prey. Interactions between a cruising predator (largemouth bass Micropterus salmoides) and an ambush predator (muskellunge Esox masquinongy) and their combination foraging on a shared prey (bluegill Lepomis macrochirus) were examined across a turbidity gradient. Turbidity modified multiple predator effects on prey. In clear water, combined predators consumed in total more prey than expected from individual predator treatments, suggesting risk enhancement for prey. In moderately turbid water, the predators consumed fewer prey together than expected, suggesting a risk reduction for prey. At high turbidity, there were no apparent emergent effects; however, the cruising predator consumed more prey than the ambush predator, suggesting an advantage for this predator. Understanding multiple predator traits across a gradient of turbidity increases our understanding of how complex natural systems function.     

Shut up or shout loudly: Predation threat and sound production in fishes

Predation is a major ecological constraint shaping behaviour and communication in animals. Several fish species are known to modify their foraging, agonistic and reproductive behaviour in the presence of predators. However, close to nothing is known about how predators affect sound production in fishes. This paper reviews how vocal fish increase their chance of survival by modifying intraspecific acoustic communication and by producing sounds directed towards predators. Field studies showed that toadfish, drums and squirrelfish reduced the number and loudness of calls when dolphin sounds were played back. These studies lack behavioural observations, leaving the question open how individual fish respond to predation threat. Croaking gouramis (Trichopsis vittata, Osphronemidae) reduced acoustic and visual signalling during dyadic contests and refrained from escalated behaviour when detecting a predator in an adjacent tank. This indicates that gouramis increase their vigilance by reducing agonistic behaviour. Vocal fish have been observed to emit sounds when predators approach or when being caught. However, none of the predators (or even conspecifics nearby) responded to these calls. Therefore, the term “predator-related” sound has been introduced in this paper to avoid implying unproven functions (alarm, startling, warning, distress and disturbance). Interestingly, the readiness of numerous taxa (e.g. catfishes) to vocalize when hand-held or netted was frequently exploited to investigate sound production in fish in relation to weight, sex, sonic organs, temperature or phylogeny. Increasing levels of noise in aquatic habitats call for more research on predator–prey interactions because of potential negative effects on detection of sounds produced by predators or prey.     

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.     

Sediment organic carbon integrates changing environmental conditions to predict benthic assemblages in shallow Arctic seas

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Non‐native rainbow trout (Oncorhynchus mykiss) occupy a different trophic niche to native Breede River redfin (Pseudobarbus burchelli) which they replace in South African headwater streams

Recent research has revealed that non‐native rainbow trout Oncorhynchus mykiss have largely replaced a native cyprinid, the Breede River redfin Pseudobarbus burchelli, as the dominant species of fish in many headwater streams in the Cape Floristic Region (CFR) of South Africa. Moreover, differences in the composition of benthic communities in CFR headwater streams with and without trout suggest that trout do not functionally compensate for the native redfin which they have replaced in these food webs. In this study, we used gut content and stable isotope analyses to characterise and compare the trophic niches and diet compositions of allopatric populations of trout and redfin in six CFR headwater streams (three containing trout, three containing redfin). Results indicate that native redfin exploit a broader trophic niche, and a more omnivorous diet, than do trout. Gut content analyses showed terrestrial invertebrates to be an important prey source for trout, which could potentially offset predation pressure on aquatic invertebrates and explain why benthic invertebrate density in streams with trout is higher than that in streams with no trout. Contrastingly, redfin diet appeared to be dominated by aquatic invertebrates, with terrestrial prey a less important food item in the guts of redfin. That redfin and trout exploit nonequivalent trophic niches may have consequences for benthic community composition in CFR headwater streams, and this study highlights the importance of quantifying how the functional role of predators changes following a predator replacement for understanding and managing the consequences of non‐native predator invasions.     

Effects of predator-induced visual and olfactory cues on 0+ perch (Perca fluviatilis L.) foraging behaviour

Abstract –  Foraging juvenile fish with relatively high food demands are usually vulnerable to various aquatic and avian predators. To compromise between foraging and antipredator activity, they need exact and reliable information about current predation risk. Among direct predator-induced cues, visual and olfactory signals are considered to be most important. Food intake rates and prey-size selectivity of laboratory-reared, naive young-of-the-year (YOY) perch, Perca fluviatilis , were studied in experiments with Daphnia magna of two size classes: 2.8 and 1.3 mm as prey and northern pike, Esox lucius , as predator. Neither total intake rate nor prey-size selectivity was modified by predator kairomones alone (water from an aquarium with a pike was pumped into the test aquaria) under daylight conditions. Visual presentation of pike reduced total food intake by perch. This effect was significantly more pronounced (synergistic) when visual and olfactory cues were presented simultaneously to foraging perch. Moreover, the combination of cues caused a significant shift in prey-size selection, expressed as a reduced proportion of large prey in the diet. Our observations demonstrate that predator-induced olfactory cues alone are less important modifiers of the feeding behaviour of naive YOY perch than visual cues under daylight conditions. However, pike odour acts as a modulatory stimulus enhancing the effects of visual cues, which trigger an innate response in perch.     

渤海鱼类群落功能群及其主要种类

根据2009年8月和10月对渤海渔业资源的底拖网调查,采用胃含物分析、聚类分析和功能群划分的方法研究了渤海鱼类群落的功能群组成及其主要种类。结果表明,渤海夏、秋季鱼类群落包括7个功能群,分别为浮游动物食性功能群、杂食性功能群、底栖动物食性功能群、虾食性功能群、虾/鱼食性功能群、鱼食性功能群和广食性功能群;其中主要功能群为浮游动物食性功能群、杂食性功能群和虾/鱼食性功能群;主要种类有小黄鱼、蓝点马鲛、斑、赤鼻棱鳀、银鲳和黄鲫。圆筛藻、中华哲水蚤、太平洋磷虾、长额刺糠虾、中国毛虾、甲壳类幼体、日本鼓虾、六丝矛尾虎鱼、小黄鱼、双壳类和腹足类是当前渤海夏、秋季鱼类群落的主要饵料种类。     

New invasive predator reduces the abundance of native prey in a cold-temperate marine fouling community

1. Invasive species threaten the conservation of marine environments, and urgent management strategies are needed to prevent their introduction, establishment and spread. Specifically, marine invasive predators destabilize ecological interactions and alter community structure and function. However, most of the evidence showing the effect of these predators is restricted to a few species and regions, while for others the information remains scarce.
2. This study experimentally evaluated the ecological effects of the sea slug Pleurobranchaea maculata, a recently introduced invasive predator, on the fouling communities of a cold-temperate port in the South-western Atlantic.
3. The results show that increasing predation pressure reduces significantly the abundance of sea anemones, which are among the preferred prey items.
4. This study reveals new evidence to help better understand the effects of P. maculata on the biological communities of a recently invaded region. Furthermore, it provides the first experimental evidence supporting the hypothesis that marine invasive predators can induce a significant decrease in the abundance of native species in benthic communities of cold-temperate marine environments in the South-western Atlantic.

     

Natural mortality augments population fluctuations of forage fish

Forage fish are a vital part of marine ecosystems, partly by supporting some of the largest fisheries worldwide, but also due to their role in food webs as prey for larger fish and other predators. One of the unresolved questions about forage fish dynamics is the causes of their significant temporal fluctuations. These fluctuations are often attributed to changes in environmental conditions, but direct correlations have proven hard to find. Here, we show how time‐varying predation mortality additionally plays a substantial role in forage fish population fluctuations. By analysing 10 stocks that have estimates of natural mortality changes through time, we find that natural mortality on average increases as population biomass declines towards a trough, and to a lesser degree decreases, when their biomass is growing towards a peak. While depensatory mortality was dominant on average in biomass dynamics leading up to peaks or troughs, some of the stocks exhibited compensatory mortality emphasizing variation between stocks. Furthermore, we show that the magnitude of natural mortality and productivity is generally higher than fishing mortality. The results underscore the importance of top‐down control on the dynamics of forage fish. We conclude that a holistic ecosystem analysis is required for a better ecological understanding of forage fish dynamics.     

莱州湾鱼类群落的营养结构及其变化

2011年5月~2012年4月期间对莱州湾进行了9次底拖网调查,对采集的20种鱼类4 854个胃含物样品进行了分析。结果表明,莱州湾鱼类群落包括浮游动物食性、底栖动物食性、鱼食性、杂食性和广食性5种食性类型,其中底栖动物食性鱼类是莱州湾各月份鱼类群落的重要食性类型,钩虾和双壳类等底栖动物饵料在莱州湾生态系统的食物关系中起着关键的作用。秋末和初春的莱州湾鱼类群落以底栖动物食性鱼类为主,食性类型单一;随着水温升高,鱼类群落的食性类型逐渐变得丰富。分析表明,莱州湾海域鱼类群落的营养结构有利于增殖放流品种后期的生长存活,建议跟踪调查增殖放流点周边海域的食物关系和饵料基础,同时结合多学科调查,选择最佳的增殖放流地点和时间,才能切实有效地保证增殖放流的效果。     

Assessing the diet of North American Atlantic salmon (Salmo salar L.) off the West Greenland coast using gut content and stable isotope analyses

Investigations on the marine feeding of Atlantic salmon (Salmo salar L.) in the Northwest Atlantic are limited compared with the Northeast Atlantic. Climate‐induced changes to food webs in Atlantic salmon feeding areas have been noted, alongside increased mortality despite a cessation of most marine fisheries. As forage efficiency may be hampering survival, it was important to address this knowledge gap. Atlantic salmon were sampled at three sites on the West Greenland coast (Sisimiut, Nuuk and Qaqortoq) between 2009 and 2011. Gut content and stable isotope analyses were combined to assess spatial and temporal differences in feeding. Capelin (Mallotus villosus) dominated the diet at Nuuk and Qaqortoq, whereas boreoatlantic armhook squid (Gonatus fabricii) was the dominant prey at Sisimiut. Hyperiid amphipods (Themisto spp.) and sand lance (Ammodytes spp.) were also important. Significant differences were found among sites for both gut contents and stable isotope analyses, with fewer differences evident temporally. Dietary differences were also evident across larger scales, with little overlap demonstrated with Northeast Atlantic diets and the emergence of boreoatlantic armhook squid as an important prey item over time. Atlantic salmon diets are frequently anchored on one or two prey items, on which they appear to specialize, but they will diversify to consume other available pelagic prey. Thus, Atlantic salmon are an opportunistic, generalist predator within the pelagic food web. The variability evident in diet suggests that the limited data available are insufficient to appropriately understand potential vulnerabilities that the species may have to ecosystem changes, and suggest further research is needed.