Reduction of Reactive Distance and Foraging Success in Smallmouth Bass, Micropterus dolomieu, Exposed to Elevated Turbidity Levels

We determined how turbidity affected the reactive distance and foraging success of smallmouth bass, Micropterus dolomieu. Smallmouth bass reactive distance decreased exponentially with increasing turbidity, from 65cm in clear water to 10cm at the highest turbidity. Turbidity significantly decreased the probability of a fish reacting to a prey item, but did not influence foraging success following reaction to the prey. Elevated turbidity may reduce stream fish foraging efficiency and decrease prey consumption.     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

Invasive carp and prey community composition disrupt trophic cascades in eutrophic ponds

The role of trophic cascades in structuring freshwater communities has been extensively studied. Most of this work, however, has been conducted in oligotrophic northern lakes that contain highly vulnerable cyprinid prey: aquatic communities where trophic interactions are likely to be stronger than in many other systems. Fewer studies have been conducted in eutrophic systems or have examined the bottom-up effects of benthivorous fishes, and none have directly compared these effects to those of piscivores on ecosystem structure and function. We conducted enclosure experiments in eutrophic ponds to examine trophic effects of invasive benthivores (common carp—Cyprinus carpio L.), native piscivores (largemouth bass—Micropterus salmoides [Lacepède]), and their interactions with common centrarchid prey with well-developed anti-predatory behaviors (age-1 bluegill—Lepomis macrochirus Rafinesque and young-of-year largemouth bass). At the end of the 60-day experiment, common carp had strong bottom-up effects that increased total phosphorus and turbidity while decreasing chlorophyll a biomass and macrophyte cover that resulted in decreased macroinvertebrate biomass and also decreased growth in both juvenile largemouth bass and bluegill. Piscivorous largemouth bass, however, did not affect the survival of either planktivorous juvenile largemouth bass or bluegill. Growth of juvenile largemouth bass was also not affected, but juvenile bluegill growth was significantly diminished, possibly due to nonconsumptive effects of predation. Our results suggest that, in a centrarchid-dominated eutrophic system, top-down effects of predators are overwhelmed by common carp-mediated bottom-up effects. These bottom-up effects strongly affected multiple trophic levels, thus altering aquatic community structure and function.     

The influence of littoral zone coarse woody habitat on home range size, spatial distribution, and feeding ecology of largemouth bass (Micropterus?salmoides)

Coarse woody habitat (CWH) may be a critical feature of lakes that influences fish distributions, movement patterns, and feeding habits. We used radio telemetry to examine the role of CWH in determining the movements of largemouth bass (Micropterus salmoides Lacepede) in the context of two whole-lake experiments that provided a gradient of four lake basins varying in natural and manipulated CWH. We also conducted diet studies on largemouth bass in these lakes to test for correlates among consumption rate and prey selectivity with bass behavior. Our results indicated that largemouth bass in basins with lower CWH abundances had larger home ranges, spent more time in deep water, were more selective predators, and showed lower consumption rates. Largemouth bass in basins with higher CWH abundances showed the opposite patterns. Low CWH abundances were correlated with a shift in largemouth bass foraging behavior from sit-and-wait to actively searching. This increased activity, coupled with the potential decline of prey fish species in the absence of CWH, may decrease largemouth bass growth potential regardless of the prey type consumed. Our results suggest that lakeshore residential development and associated removals of CWH from lakes may influence fish behavior, while CWH augmentation may reverse some of those changes. Handling editor: Steven Declerck     

Gape Limitation, Prey Size Refuges and the Top–down Impacts of Piscivorous Largemouth Bass in Shallow Pond Ecosystems

Top–down control of phytoplankton biomass through piscivorous fish manipulation has been explored in numerous ecological and biomanipulation experiments. Piscivores are gape-limited predators and it is hypothesized that the distribution of gape sizes relative to distribution of body depths of prey fish may restrict piscivore effects cascading to plankton. We examined the top–down effects of piscivorous largemouth bass on nutrients, turbidity, phytoplankton, zooplankton and fish in ponds containing fish assemblages with species representing a range of body sizes and feeding habits (western mosquitofish, bluegill, channel catfish, gizzard shad and common carp). The experimental design consisted of three replicated treatments: fishless ponds (NF), fish community without largemouth bass (FC), and fish community with largemouth bass (FCB). Turbidity, chlorophyll a, cyclopoid copepodid and copepod nauplii densities were significantly greater in FC and FCB ponds than in NF ponds. However, these response variables were not significantly different in FC and FCB ponds. The biomass and density of shallow-bodied western mosquitofish were reduced and bluegill body depths shifted toward larger size classes in the presence of largemouth bass, but the biomass and density of all other fish species and of the total fish community were unaffected by the presence of largemouth bass. Our results show that top–down impacts of largemouth bass in ecosystems containing small- and deep-bodied fish species may be most intense at the top of the food web and alter the size distribution and species composition of the fish community. However, these top–down effects may not cascade to the level of the plankton when large-bodied benthivorous fish species are abundant.     

Susceptibility of Transgenic and Wildtype Zebra Danios, Danio rerio, to Predation

The effects to ecosystems by genetically modified organisms are still unknown, yet a transgenic version (the red glofish or red fluorescent protein (RFP) transgenic zebra danio) of the zebra danio, Danio rerio, a common aquarium fish, has become the first transgenic pet sold in the USA. It has been hypothesized that RFP zebra danios will not persist in nature because they will be preferentially preyed upon due to their red coloration; however, the bright coloration of wildtype zebra danios may indicate they are aposematic since they are not preyed upon immediately by predators in their native range. These hypotheses were addressed via nine predation experiments with largemouth bass, Micropterus salmoides, as predator and combinations of mosquitofish, Gambusia affinis, wildtype zebra danios, and RFP zebra danios as prey. Neither wildtype nor RFP transgenic zebra danios are aposematic since both varieties were readily consumed by largemouth bass in laboratory trials. Both varieties were preyed upon in approximately equal proportion (1.4 to 1.0) so the bright, apparently conspicuous coloration of the transgenic zebra danios did not increase their susceptibility to predation. In these laboratory trials, largemouth bass did not preferentially prey upon a native fish, the mosquitofish, relative to wildtype zebra danios (1.2 to 1.0). Based on the results of these experiments, wildtype zebra danios and RFP transgenic zebra danios are likely to be preyed upon in a similar fashion as native forage fish.     

Characteristics of reaction field and the reactive distance of a planktivore, Pseudorasbora parva (Cyprinidae), in various environmental conditions

The characteristics of the reaction field and the reactive distance of the Stone moroko (Pseudorasbora parva) were studied under three environmental conditions (structural complexity, light intensity and turbidity) and three prey sizes. In optimal experimental conditions, under no structural complexity, light intensity of 200 lux and turbidity less than 1 NTU (Nephelometric Turbidity Units), the cross-section of the reaction field was found to be elliptic with a bearing angle larger than the elevation angle, but both angles changed slightly depending on environmental conditions. The reactive distance was large, and the fish frequently attacked prey that was located within 15–60 degrees to each side from the frontal direction of a fish (i.e., ± 15 degrees from the axis of the fish body) horizontally. In the light intensity below 50 lux or turbidity above 10 NTU, however, the attack frequency and the reactive distance in the frontal direction of a fish did not differ from other horizontal directions in the reaction field. The average reactive distance increased proportionally with increasing strand distance, but it gradually reached a constant value for strand distances greater than about 3.6 times the fish body length. The average reactive distance increased in the light intensity range of 10–200 lux and decreased negatively with turbidity increasing. The average reactive distance increased with larger prey size, but the rate of increase of the reactive distance gradually decreased.     

Effects of turbidity and an invasive waterweed on predation by introduced largemouth bass

Anthropogenic activities lead to changes in characteristics of aquatic ecosystems, including alteration of turbidity and addition of invasive species. In this study, we tested how changes in turbidity and the recent invasion of an aquatic macrophyte, Egeria densa, may have changed the predation pressure by introduced largemouth bass on juvenile striped bass and delta smelt, two species that have seen a drastic decline in recent decades in the Sacramento-San Joaquin Delta. In a series of mesocosm experiments, we showed that increases in vegetation density decreased the predation success of largemouth bass. When placed in an environment with both open water and vegetated areas, and given a choice to forage on prey associated with either of these habitats, largemouth bass preyed mainly on open water species as opposed to vegetation-associated species, such as juvenile largemouth bass, bluegill or red swamp crayfish. Finally, we showed that turbidity served as cover to open water species and increased the survival of delta smelt, an endemic species at risk. We also found that such open water prey tend not to seek refuge in the vegetation cover, even in the presence of an imminent predation threat. These results provide the beginning of a mechanistic framework to explain how decreases in turbidity and increases in vegetation cover correlate with a decline of open water species in the Sacramento-San Joaquin Delta.     

Antipredator Behavior of American Bullfrogs (Lithobates catesbeianus) in a Novel Environment

Invasive species capable of recognizing potential predators may have increased establishment rates in novel environments. Individuals may retain historical predator recognition and invoke innate responses in the presence of taxonomically or ecologically similar predators, generalize antipredator responses, or learn to avoid risky species in novel environments. Invasive amphibians in aquatic environments often use chemical cues to assess predation risk and learn to avoid novel predators via direct experience and/or associated chemical cues. Ontogeny may also influence recognition; experience with predators may need to occur at certain developmental stages for individuals to respond correctly. We tested predator recognition in invasive American bullfrog ( Lithobates catesbeianus) tadpoles that varied in experience with fish predators at the population and individual scale. We found that bullfrog tadpoles responded to a historical predator, largemouth bass ( Micropterus salmoides), only if the population was locally sympatric with largemouth bass. Individuals from a population that did not co‐occur with largemouth bass did not increase refuge use in response to either largemouth bass chemical cues alone or chemical cues with diet cues (largemouth bass fed bullfrog tadpoles). To test whether this behavioral response was generalized across fish predators, we exposed tadpoles to rainbow trout ( Oncorhynchus mykiss) and found that tadpoles could not recognize this novel predator regardless of co‐occurrence with other fish species. These results suggest that environment may be more important for predator recognition than evolutionary history for this invasive species, and individuals do not retain predator recognition or generalize across fish predators.     

Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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Growth of pike larvae: effects of prey, turbidity and food quality

We studied experimentally the effects of turbidity and prey composition on pike larval growth and hypothesized that pike larval growth varies with turbidity and food quality. We reared the first-feeding pike larvae (Esox lucius) in laboratory tanks with (1) clear or (2) turbid water provided with zooplankton rations from (3) an inner and (4) an outer archipelago site. The sites differ in physical features, salinity, eutrophication status, zooplankton community structure and density. Pike larvae showed the highest weight increase in clear water with zooplankton from the outer site and the poorest weight increase in turbid water with zooplankton as prey from the inner site. Our fatty acid analysis revealed that unsaturated fatty acid levels were highest in the outer site. The relative percentage of copepods was also higher in the outer site. This study supports the hypothesis that turbidity weakens the ability of pike larvae to capture certain prey. Further, zooplankton community composition matters in turbid water, but is not a primary factor in clear water.     

Relative effects of turbidity and light intensity on reactive distance and feeding of an estuarine fish

Synopsis Gulf killifish Fundulus grandis were allowed to prey on daggerblade grass shrimp Palaemonetes pugio in clear water with bright light, turbid water containing bentonite clay, and clear water treatments where the light intensity was adjusted to match that in the bottom of the turbid tanks. Significantly fewer shrimp were consumed in the turbid tanks relative to the clear and shade treatments where predation rates did not differ significantly. The results suggested that the influence of suspended particles on predation rates was a consequence of light scattering and was not related to a decrease in light intensity. Reactive distances were subsequently determined for human observers viewing a small target in elongated tanks containing turbid water (7.3–60.5 NTU) under conditions of both low (8-10 E m^–2 s^–1) and high illumination (153–1249 E M^–2 s^–1). Reactive distance was primarily governed by turbidity while light intensity had little influence except at low turbidities. The shape of the relationship between reactive distance and turbidity for humans resembled curves reported for a variety of fish species.     

Growth,survival, and foraging abilities of early life history stages of blue tilapia,Oreochromis aureus,and largemouth bass,Micropterus salmoides

Synopsis I compared growth, survival, and foraging abilities of early life history stages of blue tilapia,Oreochromis (=Sarotherodon = Tilapia) aureus, and largemouth bass,Micropterus salmoides, at equivalent prey (zooplankton) densities in the laboratory. Survival and growth of blue tilapia exceeded those of largemouth bass and, at comparable ages, blue tilapia were superior to largemouth bass in ability to prey on zooplankton; the enhanced survival and growth of blue tilapia appeared to result from their larger initial (egg) size. Predatory abilities of the species at equivalent sizes were similar. The success of blue tilapia in colonizing new systems and rapidly achieving high abundances in Florida may be a consequence of large initial size.     

The energetic impact of overwinter prey assemblages on age-0 largemouth bass, <Emphasis Type="Italic">Micropterus salmoides</Emphasis>

Synopsis We compared survival, growth, and swimming performance of two size classes of age-0 largemouth bass, Micropterus salmoides, in the spring after being fed diets of bluegill, Lepomis macrochirus, fathead minnows, Pimephales promelas, or invertebrate prey during the winter. Regardless of prey assemblage, survival was uniformly high and independent of size. Length, wet- and dry-mass, and condition was also similar among treatments for both size classes. However, variation in individual performance differed, with the lowest variability in growth occurring among small age-0 largemouth bass in the invertebrate only treatment. Absolute and length corrected swimming speeds of largemouth bass were highest for invertebrate prey assemblages, intermediate for fathead minnow prey, and lowest for bluegill prey. The patterns in growth and spring swimming performance likely reflect the varied nutritive quality of different prey, the ability of largemouth bass to capture different prey, and competition with the piscine prey.     

Algal Turbidity Reduces Risk Assessment Ability of the Three‐Spined Stickleback

Recent anthropogenic increases in algal turbidity in aquatic habitats have been suggested to affect the ability of fish to assess predation risk. We investigated the response of feeding three‐spined sticklebacks (Gasterosteus aculeatus) exposed to a sudden appearance of an avian predator (the silhouette of common tern, Sterna hirundo), under clear and turbid water conditions. As stickleback use social cues to aid in predator avoidance, we also tested whether turbidity affected social information use by manipulating group size. We found that in turbid water, a smaller proportion of fish would escape from the feeding area, that the distance escaped was shorter and that a smaller proportion of fish fled into shelter. Larger group size was associated with longer escape distance and greater shelter use. However, there was no effect of group size on the proportion of fish that escaped the arena. The effect of group size was similar for turbid and clear water. Our finding that the fish showed a weaker antipredator response suggests that turbidity impedes their risk assessment capability. However, the sticklebacks were still able to benefit of the social facilitation provided by being in a group. This suggests that algal turbidity has detrimental effects on the ability of sticklebacks to assess predation risk from avian predators in shallow water. An implication is that in shallow water fish may be more vulnerable to avian predation under turbid conditions.     

Turbidity decreases anti-predator behaviour in pike larvae, <Emphasis Type="Italic">Esox lucius</Emphasis>

Synopsis We tested how algal turbidity and light conditions influence anti-predator behaviour of first-feeding pike. Results showed that pike larvae were able to detect the predator by both chemical and visual signals in turbid water. However, the anti-predator behaviour was reduced in turbid water compared with clear water. Larvae hid in the vegetation in the presence of predator cues more in clear water than in turbid water. The attack rate on zooplankton in clear water was lower in the presence of predator cues, whereas no such difference was detected in turbid water. Both of these results indicate that turbidity acted as a refuge for larvae. Light proved to be an important regulating factor for feeding pike in the absence of predators, demonstrated as lowered attack rates in 50 light level in both clear and turbid water. This indicates that long-term turbidity may be critical for small larvae, which need to feed continuously to survive.     

Interplay of individual interactions and turbidity affects the functional response of three-spined sticklebacks Gasterosteus aculeatus

The effects of turbidity, size and the presence of conspecifics on the functional response, feeding latency and activity in the three-spined stickleback Gasterosteus aculeatus were examined. A significant interaction between standard length and presence of conspecifics demonstrated an increase in attack rates of larger individuals in the presence of conspecifics. Attack rate was also higher in turbid water. Feeding latency decreased with prey concentration and presence of conspecifics, but was not affected by turbidity. Activity level did not change with prey levels, but increased with turbidity. These results can help to better understand how individual flexibility in the functional response can affect prey mortality according to environmental perturbation and social interaction at the level of the predator.     

Effects of turbidity and risk of predation on habitat selection decisions by Fathead Minnow (<Emphasis Type="Italic">Pimephales promelas</Emphasis>)

Within aquatic ecosystems, turbid environments will have a significant impact upon predator-prey interactions if both the predator and their prey rely upon vision as their primary sense. Increasing water turbidity will reduce the probability of being detected by a predator, and once detected should provide prey with cover that is close and ubiquitous. We tested the extent that these features of a turbid environment will have in affecting the impact of predation risk on habitat quality using Fathead Minnow (Pimephales promelas) as the prey, and Yellow Perch (Perca flavescens) and Black Bullhead (Ameiurus melas) as visual and non-visual predators, respectively. Our experiments demonstrated a strong preference for turbid habitats in the absence of a predator. When a predator was present in a turbid habitat, the minnows reduced their use of this location but still preferred it to a clear habitat with no predator. These data suggest turbidity confers a benefit to feeding Fathead Minnow that more than compensates for the cost of predation risk.     

Performance as a fish predator of largemouth bass [<Emphasis Type="Italic">Micropterus salmoides</Emphasis> (Lacepède)] invading Japanese freshwaters: a review

This review focuses on how predator performance of the invasive largemouth bass [Micropterus salmoides (Lacepède)] has been, or will be, formed in Japanese freshwaters. Predation impacts of largemouth bass on fish communities appear pervasive in both Japanese as well as North American freshwaters. Factors affecting performance as a piscivorous predator are (1) light intensity and water clarity, (2) oxygen depletion, (3) prey size and gape size, (4) behavioral refuge of prey, (5) weed beds as refuge for prey fish, (6) interaction with bluegill. Size and behavioral refuges requirements are so rigorous that they may have evolved only in some North American prey fish species like bluegill; therefore, most Japanese native fish species are unlikely to be equipped with such refuges. However, refuge habitats like aquatic weed beds could develop in Japanese freshwaters, allowing prey fish species to survive under predation pressure. The density, architecture, and species composition of aquatic plants may affect their suitability as refuges. Studies in Japanese waters have suggested that the presence of rich aquatic vegetation or invasive bluegill in bass-introduced waters have suppressed the predation impact of largemouth bass on fish communities. In addition to these environmental factors, original genotypic and phenotypic traits of the introduced largemouth bass, and hybridization between different lineages of largemouth bass or with Florida bass [Micropterus floridanus (Lesueur)] may be involved in further adaptation of invasive largemouth bass to Japanese freshwaters.